General Animal Nutrition

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Animal nutrition
Kroly Doublecz

Animal nutrition
Kroly Doublecz
Publication date 2011

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Table of Contents
Cover .................................................................................................................................................. x
1. Prediction the worlds potential population, prospects for food and water .................................... 1
1. .............................................................................................................................................. 1
2. Prospects for water and food .......................................................................................................... 2
1. Food in particular and arable products in general ................................................................. 3
2. Livestock production ............................................................................................................. 4
3. Fishing ................................................................................................................................... 4
4. The animal and its food ......................................................................................................... 5
5. Water ..................................................................................................................................... 5
3. Analysis of foods ............................................................................................................................ 8
1. Proximate analysis of foods .................................................................................................. 8
2. Modern analytical methods ................................................................................................... 8
3. Starch and sugars .................................................................................................................. 9
4. Fibre ...................................................................................................................................... 9
5. Minerals .............................................................................................................................. 10
6. Amino acids, fatty acids and sugars .................................................................................... 10
7. Measurement of protein in foods for ruminants .................................................................. 11
8. Spectroscopy ....................................................................................................................... 11
4. The significance of antinutritional factors .................................................................................... 12
1. Lectins ................................................................................................................................. 12
2. Protease inhibitors ............................................................................................................... 13
3. Alpha-amylase inhibitor ...................................................................................................... 14
4. Tannins ................................................................................................................................ 14
5. Flatulence factors ................................................................................................................ 15
6. Antigenic proteins ............................................................................................................... 16
7. Phytic acid ........................................................................................................................... 16
8. Vicine and convicine ........................................................................................................... 16
9. Saponins .............................................................................................................................. 16
10. Glucosinolates ................................................................................................................... 17
11. Oxalic acid ........................................................................................................................ 17
12. Gossypol ........................................................................................................................... 17
13. Alkaloids ........................................................................................................................... 17
14. Sinapins ............................................................................................................................. 18
15. Non-starch polysaccharides ............................................................................................... 18
16. Mechanisms by which dietary NSPs affect animal performance ...................................... 19
5. Feed additives ............................................................................................................................... 21
1. Antibiotics ........................................................................................................................... 21
2. Modes of action antibiotics ................................................................................................. 22
3. Probiotics ............................................................................................................................ 22
4. Prebiotics ............................................................................................................................. 23
5. Enzymes .............................................................................................................................. 24
6. Enzyme action in pigs and poultry ...................................................................................... 25
7. Organic acids ....................................................................................................................... 26
8. Antimicrobial action of organic acids ................................................................................. 27
9. Modifiers of rumen fermentation ........................................................................................ 27
6. Advances in feed evaluation for pigs ............................................................................................ 29
1. Variability in feeding value within the same feed ingredient .............................................. 30
2. Effects of processing on the feeding value of feed ingredients ........................................... 31
3. Interactive effects of feed ingredients on nutrient digestion and utilisation ........................ 31
4. Significance of microbial fermentation in the upper gut of pigs ......................................... 32
5. Feed ingredient specific effects on nutrient metabolism in visceral organs ........................ 33
6. Differences in nutrient digestibilities between different classes of pig ............................... 34
7. Advances in feed evaluation for poultry ....................................................................................... 36
1. Feed evaluati ....................................................................................................................... 36
2. Energy ................................................................................................................................. 36
3. Experimental methods in feed energy evaluation ............................................................... 36

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4. Prediction of the ME values of poultry feeds and feedstuffs .............................................. 38
5. Prediction equations for feedstuffs ...................................................................................... 38
6. Prediction equations for feed fats ........................................................................................ 38
7. Interactions among energy yielding ingredients .................................................................. 39
8. Enhancing dietary energy values ......................................................................................... 39
9. Feed enzymes ...................................................................................................................... 39
10. Protein and amino acids .................................................................................................... 40
11. Measurement of amino acid availability ........................................................................... 40
12. Feed formulation and the control of protein and amino acid levels .................................. 40
13. Information sources for amino acids and protein digestibility .......................................... 40
14. Minerals ............................................................................................................................ 41
8. Advances in feed evaluation for ruminants .................................................................................. 42
1. Energy evaluation systems .................................................................................................. 42
1.1. Early energy systems .............................................................................................. 42
1.2. Recent systems ....................................................................................................... 42
2. Protein evaluation for ruminant animals ............................................................................. 44
3. Degradability of the nitrogen fraction of the diet ................................................................ 44
3.1. Measurement of degradability in vivo .................................................................... 45
3.2. Determination of degradability in sacco (or in situ) ............................................... 45
4. Laboratory procedures for determining nitrogen degradability .......................................... 46
4.1. Solubility in buffer solutions .................................................................................. 46
4.2. Solubility in enzyme solutions ................................................................................ 46
4.3. Near-infrared reflectance spectroscopy (NIRS) ...................................................... 46
4.4. Rate of passage ....................................................................................................... 46
4.5. Efficiency of nitrogen capture ................................................................................ 47
4.6. Yield of microbial protein ...................................................................................... 47
4.7. True digestibility of protein .................................................................................... 47
4.8. Efficiency of utilisation of absorbed amino acids ................................................... 48
4.9. The Hungarian metabolisable protein system ......................................................... 49
5. Questions: ........................................................................................................................... 49
9. Advances in feed evaluation for companion animals ................................................................... 50
1. Anatomy of the digestive tract of cats and dogs .................................................................. 50
2. Metabolic adaptations of the cat and the dog ...................................................................... 50
2.1. Protein and amino acid metabolism ........................................................................ 50
2.2. Essential fatty acids ................................................................................................ 51
2.3. Vitamin metabolism ............................................................................................... 51
2.4. Mineral metabolism ................................................................................................ 51
3. Criteria for diet formulation ................................................................................................ 52
3.1. Palatability testing .................................................................................................. 52
3.2. Tests for nutritionally complete and balanced diets: chemical testing .................... 53
3.3. Other nutritional tests for companion animal diets ................................................. 53
4. Prepared cat and dog foods ................................................................................................. 53
5. Questions: ........................................................................................................................... 54
10. Recent advances in feed formulation .......................................................................................... 55
1. Growth and production characteristics of the livestock ...................................................... 55
2. Tissue nutrient requirements to meet production goals ....................................................... 56
3. Translation of tissue requirements into dietary specifications ............................................ 56
4. Nutrient composition of feed ingredients ............................................................................ 57
5. Minimum and maximum inclusion limits for nutrients and ingredients ............................. 57
6. Formulation of diets to least-cost ........................................................................................ 58
7. Manufacture of the feed ...................................................................................................... 59
8. Questions: 1. What kind ...................................................................................................... 59
11. Animal nutrition and the consumers of animal products ............................................................ 61
1. The contribution of animal products to human requirements .............................................. 61
2. Ethical and environmental objections to the use of animal products .................................. 63
3. Nutrition and human health ................................................................................................. 64
4. Future trends in the consumption of animal products ......................................................... 66
5. Questions: ........................................................................................................................... 66
12. Safety and quality of food from animals ..................................................................................... 67
1. Food charter ........................................................................................................................ 67

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1.1. Independence .......................................................................................................... 67
1.2. Safety ...................................................................................................................... 67
1.3. Quality .................................................................................................................... 67
1.4. Choice ..................................................................................................................... 67
1.5. Information ............................................................................................................. 67
1.6. Redress ................................................................................................................... 67
1.7. Accountability ........................................................................................................ 67
1.8. Representation ........................................................................................................ 68
2. Representation ..................................................................................................................... 68
2.1. Risk ......................................................................................................................... 68
2.2. Safety ...................................................................................................................... 69
2.3. Nutrition ................................................................................................................. 69
2.4. Composition ........................................................................................................... 69
2.5. Function .................................................................................................................. 69
2.6. Sensory perceptions ................................................................................................ 69
2.7. Authenticity ............................................................................................................ 70
2.8. Convenience ........................................................................................................... 70
2.9. Exotic, novel, luxury .............................................................................................. 70
2.10. Labelling and packaging ....................................................................................... 70
3. Consumer awareness ........................................................................................................... 70
4. Questions: ........................................................................................................................... 70
13. Environmental impacts of feeding monogastric animals ............................................................ 72
1. Nutrition and mineral excretion .......................................................................................... 72
2. Nutrition and heavy metals ................................................................................................. 73
3. Nutrition and ammonia emission ........................................................................................ 73
3.1. Lowering crude protein intake ................................................................................ 73
3.2. Shifting nitrogen excretion from urea in urine to protein in faeces ........................ 73
3.3. Lowering the pH of faeces ...................................................................................... 74
3.4. Lowering the pH of urine ....................................................................................... 74
4. Nutrition and odour emission .............................................................................................. 74
5. Nutrition and methane emission .......................................................................................... 75
6. Questions: ........................................................................................................................... 75
14. Environmental impacts of feeding ruminants ............................................................................. 76
1. Environmental pollution caused by animal production ....................................................... 76
2. Feeding Strategies and Excretion of Waste ......................................................................... 77
2.1. Energy Losses ......................................................................................................... 77
2.2. Nitrogen Losses ...................................................................................................... 78
2.3. Phosphorus Losses .................................................................................................. 79
2.4. Potassium losses ..................................................................................................... 80
3. Implications ......................................................................................................................... 81
4. Questions: ........................................................................................................................... 81
15. Nutritional evaluation of the first generation genetically modified plants (GMP) ..................... 82
1. Studies with Bt (Bacillus thuringiensis)-maize ................................................................... 82
1.1. Beef cattle ............................................................................................................... 82
1.2. Sheep ...................................................................................................................... 83
1.3. Growing and, finishing pigs ................................................................................... 83
1.4. Laying hens ............................................................................................................ 84
1.5. Broilers ................................................................................................................... 85
2. Bt-potatoes .......................................................................................................................... 85
3. Glufosinate tolerant (Pat) maize in pigs .............................................................................. 86
4. Glufosinate tolerant (Pat) sugar beets ................................................................................. 88
4.1. Sheep ...................................................................................................................... 88
4.2. Pigs ......................................................................................................................... 88
5. Roundup ready (RR, Glyphosate tol.) soybeans in pigs ...................................................... 89
6. Summary ............................................................................................................................. 89
7. Questions: ........................................................................................................................... 90
16. Nutritional evaluation of the second generation genetically modified plants (GMP) ................. 91
1. Increased myristic and palmitic acid in rapeseed for pigs ................................................... 91
2. Inulin synthesizing potatoes in pigs .................................................................................... 93
3. Fate of DNA ........................................................................................................................ 93

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4. Questions: ........................................................................................................................... 94
17. References .................................................................................................................................. 95

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List of Figures
2.1. Table 1. Population and water resources by major world regions (Cole, 1999) .......................... 2
2.2. Table 2. Cropland in millions of hectares and hectares per capita (Cole, 1999) .......................... 3
2.3. Fig. 1. The main components of foods, plants and animals ......................................................... 5
2.4. Table 3. Composition of some plant and animal products (g/kg) ................................................ 6
3.1. Table 4. The fibre components of some common foods (g/kg D.M.) .......................................... 9
4.1. Table 5. Effects of feeding soya bean lectins on flows of dry matter and nitrogen at the distal ileum in
growing pigs (Schulze et al., 1995) .................................................................................................. 13
4.2. Table 6. Effcet of alternative heat treatments on lectin and trypsin inhibitor activity in phaseolus
beans (van der Poel, 1999) ................................................................................................................ 14
4.3. Table 7. Contents of antinutritional factors and crude protein digestibility values of raw and
germinated beans, measured with growing pigs ............................................................................... 14
4.4. Table 8. Apparent ileal and faecal digestibilities of crude protein and nitrogen free extract in different
cultivars of faba beans to piglets, included at 300g/kg (Jansmann, 1993) ........................................ 15
4.5. Figure 2. Polysaccharide structures commonly found in feed ingredients of plant origin (de Lange,
2000) ................................................................................................................................................. 18
5.1. Fig. 3. Competitive exclusion of pathogen bacteria due to attachment of non-pathogens ......... 23
5.2. Fig. 4. Effect of oligosaccharides on the excretion of harmful bacteria .................................... 24
5.3. Table 9. Effect of -glucanase addition to barley on the production parameters of 3 week old broiler
chicks ................................................................................................................................................ 25
5.4. Table 10. Effect of graded levels of fumaric acid on the growth rate and food conversion efficiency of
young pigs. Easter R.A. (1988): Acidification of diets for pigs, In: Haresign, W. and Cole D.J.A. Recent
Advances in Animal Nutrition .......................................................................................................... 26
6.1. Table 11. Determined and calculated amino acid digestibilities for selected amino acids ........ 32
6.2. Table 12. Distribution of NSP fermentation in the gastrointestinal tract of the pig (Graham s mtsai.,
1986; Longland s mtsai., 1990; Zhu s mtsai., 1993; Millard s Cheeson,1992 ............................. 33
6.3. Table 13. Apparent ileal digestibilities of amino acids in growing pigs and lactating sows (de Lange
and Fuller, 2000) ............................................................................................................................... 34
7.1. Figure 5. The relationship between feed intake and energy excretion and different ME values (Fisher,
2000) ................................................................................................................................................. 37
8.1. Fig 6. Relationship of protein disappearance to time of incubation ........................................... 45
8.2. Table 14. Composition of rumen bacteria (g/kg dry matter) ..................................................... 47
8.3. Table 15. Amino acid composition of ruminal bacteria (g/100g of amino acids) ...................... 48
8.4. Table 16. Amino acid composition of whole and insoluble protein in some common foods (g/100g
protein) ............................................................................................................................................. 49
9.1. Table 17. Minimum dietary crude protein and total essential amino acid requirements for cats, dogs
and rats (Hendriks, 1996) ................................................................................................................. 51
9.2. Fig. 7. True ileal digestibility of amino acids and amino acid nitrogen in moist canned cat food after
different heating periods (Hendriks et al., 1999) .............................................................................. 54
10.1. Fig. 8. Comparison the results of the cost/tonne and cost/MJ DE formulations ...................... 58
10.2. Table 18. Effect of the form of feed on the fed wastage and performance of pigs (Hutson, 1998)
59
11.1. Table 19. Contribution of various food groups to world food supplies (FAO) ........................ 61
11.2. Table 20. Meat and milk consumption in selected countries and world regions (kg/head/year) (FAO)
........................................................................................................................................................... 61
11.3. Table 21. Contribution of animal products to human diets (FAO, 1998) ................................ 61
11.4. Table 22. Some important diseases transmissible in food from farm animals to man ............. 64
12.1. Fig. 9. Food quality: the consumer priorities ........................................................................... 69
13.1. Fig. 10. Relationship between NSP content of the diet and the urine-Ni/faecal-N ratio (Jongbloed,
2001) ................................................................................................................................................. 74
13.2. Table 23. Overview of the volatile products formed by microbial activity in manure from the main
components in urine and faeces ........................................................................................................ 74
14.1. Fig. 11. The effect of annual milk yield and number of lactations on the input of fossil energy (FE)
per kilogram of milk. ........................................................................................................................ 77
15.1. Table 24. Fattening and slaughter results of bulls (n=20) (Aulrich et al., 2001) ..................... 82
15.2. Table 25. Digestibility coefficients and metabolisable energy content of Bt-maize silage in sheep as
compared to that of the isogenic line (Aulrich et al., 2001) .............................................................. 83

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15.3. Table 26. Chemical composition of transgenic maize seeds as compared to that of the parental line
(Reuter et al., 2002) .......................................................................................................................... 83
15.4. Table 27. Coefficient of digestibility and energetic feeding value of maize for pigs (Reuter et al.,
2002) ................................................................................................................................................. 83
15.5. Table 28. Growth performance of growing-finishing pigs fed iso- or transgenetic maize diets over a
period of 91 days (Reuter et al., 2002) ............................................................................................. 84
15.6. Table 29. Chemical composition of Bt-maize seeds and the isogenic comparator used in the trials
with poultry (Aulrich et al., 2001) .................................................................................................... 84
15.7. Table 31. Performance of broilers fed Bt-maize or the isogenic comparator as the principal
component in the diet (Aulrich et al., 2001) ..................................................................................... 85
15.8. Table 32. The influence of non-GM and GM potatoes on feed intake, final body weight and feed
conversion of male broilers (from days 14 ton28 of age) (Halle et al., 2005) .................................. 86
15.9. Table 33. Proximate analysis, starch, sugar and NSP-composition of Pat-maize seeds compared with
those of the corresponding non-transgenic lines (g/kg DM) (Bhme et al., 2001) ........................... 86
15.10. Table 34. Amino acid analysis of Pat-maize seeds compared with the corresponding non-transgenic
controls (amino acids g/16g N) (Bhme et al., 2001) ....................................................................... 86
15.11. Table 35. Fatty acid composition of Pat-maize grains compared with the corresponding transgenic
controls (percent of total fatty acids) (Bhme et al., 2001) .............................................................. 87
15.12. Table 36. Coefficient of digestibility and energy content of Pat-maize grains for pigs as compared
with those of the non-transgenic control (Bhme et al., 2001) ......................................................... 87
15.13. Table 37. Proximate analysis and sugar contents of Pat-sugar-beets and Pat-sugar-beet top silage as
compared to those of the corresponding non-transgenic line (g/kg of DM) (Bhme et al., 2001) ... 88
15.14. Table 38. Digestibility coefficient and eneryg content of Pat-sugar beets for pigs as compared with
those of the non-transgenic control (Bhme et al., 2001) ................................................................. 88
15.15. Table 39. Composition (g/kg DM) of iso- and transgenic full-fat roasted soybeans fed to growing-
finishing pigs (Flachowsky et al., 2007) ........................................................................................... 89
15.16. Table 40. Performance of pigs over 42 days of feeding grower-finisher diets containing isogenic or
Roundup Ready full-fat roasted soybeans (Flachowsky et al., 2007) ............................................... 89
16.1. Table 41. Chemical composition of iso- and transgenetic rapeseed ........................................ 91
16.2. Table 42. Coefficient of digestibility and energy content of rapeseed-based diets and performance
parameters of pigs (n=10, 32-105 kg body weight) .......................................................................... 92
16.3. Table 43. Selected proximate analysis, starch, macro-elements, amino acids and glycoalkaloids of
transgenic inulin synthesising potatoes compared with those of the parenteral line ......................... 93
16.4. Table 44. Studies of the transfer of foreign DNA fragments into farm animals .................. 94

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List of Tables
1. ........................................................................................................................................................ x

x
Cover
Animal nutrition
Author:
Kroly Dublecz
Az Agrrmrnki MSc szak tananyagfejlesztse
TMOP-4.1.2-08/1/A-2009-0010 projekt
Table 1.

1
Chapter 1. Prediction the worlds
potential population, prospects for
food and water
1.
It is possible to make projections for various aspects of world production and consumption, indicating both total
amounts and amounts per capita. What the natural resource or item of production or consumption considered
(e.g. area under forest, natural gas production, passenger cars in circulation) the absolute amounts can be given
and the per capita amounts are then calculated.
In order to make immediate comparisons between various items possible, 1990 always has an index of 100,
whatever the absolute quantity involved. Three alternative futures are then projected to the year 2050. In each
case, two of these are always the same. In one projection the 1990 total stays unchanged to the year 2050, in
which case the per capita level drops to about 55 per cent between 1990 and 2050 on account of expected
population growth. In another projection, the per capita level stays the game and the total amount therefore has
to increase by more than 85 per cent to allow for the expected increase in population. A third alternative
projection, calculated is simply the continuation of the 1950-1990 trend.
The population of the world will probably reach soon 7 billion and will therefore have grown almost four times
in the 20th century, a cause for alarm or satisfaction, depending on one's views. At the same time, the total
consumption of non-renewable natural resources is at least ten times as high in the 1990s as it was in the 1890s.
Population is changing at very different rates in different regions and countries of the world. In some countries
of Central Europe it is now slowly declining, whereas on average it is increasing by about 2.6 per cent annually
in Africa, a 'compound interest' rate that would result in a doubling of population in 26 years. In Europe and
North America the reduction of fertility in recent decades has largely been spontaneous whereas in China
draconian measures have been applied since the 1970s to achieve as many one-child families as possible. In
India birth control methods have been less drastic and less effective.
Since population change is determined by tens of millions of 'decisions' each year to have children it is unlikely
that anything other than a devastating natural or man-made disaster could suddenly change the present trend. A
total world population of around 10 billion is anticipated by the author for 2050, a figure that falls within the
range of several other projections. There will be a much smaller share of the total population in the present
developed countries in 2050 than now, a situation affected in only a very limited way by international migration.
In 2050 the global demographic picture will be characterised by two outstanding features: the presence of a very
large rural, predominantly agricultural, population on a correspondingly very limited area of farmland, and the
presence of numerous very large cities, many of them the capitals of larger developing countries.

2
Chapter 2. Prospects for water and
food
Water
The need for water seems likely to increase with the growth of the world population expected in the next 50-60
years. Even if global warming modifies the climate by 2050 it is unlikely that the broad picture of the
distribution of fresh water would change greatly. Therefore the existing water sources have to be used more
intensively. Three futures for water can be contrasted:
1., The population of the world increases to only 9 billion in 2050. Extensive transfers of fresh water by canal or
pipeline move water into areas where cultivation is at present precluded or yields are low through lack of
precipitation: for example, Uzbekistan, parts of western USA, northern China. Water supply in growing urban
areas in developing countries is improved. Water use, especially for domestic purposes, is reduced in developed
countries, possibly through some form of rationing, such as a surcharge on water consumed over a certain level.
Desalination is practised on a much larger scale than now. While such developments would improve water
supply in various parts of the world in the next few decades, the environment could be adversely affected, both
by construction works and by emissions from the fossil fuels used to desalinate water, at least until enough solar
power or power from nuclear fusion becomes widely available.
2., Recent trends continue.
3., The population of the world increases to around 11 billion. Little is done to facilitate new transfers of water,
to use irrigation water more efficiently, or to economise on water use in developed countries. Aquifers are used
up more quickly than at replacement rate, a widespread feature now in India.
The issue of water supply will be crucial already in the in the first half of the 21st century. Table 1. shows the
mounting pressure on water resources in the developing regions due to population growth. It shows the effect of
expected population growth between 1991 and 2025 on per capita resources, particularly in all of Africa and in
Southwest Asia, the regions in which the fastest population growth in the world is currently being experienced
and where it seems likely to continue
Figure 2.1. Table 1. Population and water resources by major world regions (Cole,
1999)

Prospects for water and food

3
Notes: 1 China, Japan and Southeast Asia 2 Excluding former USSR countries 3 Includes Mexico and Central
America
Since irrigation for agricultural purposes is by far the largest user of water in many countries Table 2. has been
compiled to show the countries with the largest areas of irrigated land in the world. In China about half of the
cultivated land is irrigated, in India almost a third. In several smaller countries the proportion is much higher, as
for example in Egypt, where it is almost 100 per cent. The worlds twelve largest users of water for irrigation
account for 70 per cent of all water consumed for irrigation, while Asia uses about 64 per cent of the total.
Figure 2.2. Table 2. Cropland in millions of hectares and hectares per capita (Cole,
1999)

Some form of irrigation is practised in many countries of the world, but there is still the potential in many places
both to make greater use of water from rivers, and to use the present supply more efficiently.
1. Food in particular and arable products in general
Different prospects for food production are so described only with reference to food since a future that may be
favourable for food production may be unfavourable for other users of the land. From different combinations the
optimistic scenario would be that the number of people to be fed would increase only by 50 per cent, an increase
in the area under cultivation of 70-80 per cent, a widespread increase in yields, and an organisation in place to
ensure flexible transfers of food as and when needed.
The worst forecast mean a future would result in a desperate food situation by 2050, with an 80-90 per cent
increase in the number of people to be fed, rather less land to cultivate, little change in yields, and each major
region or country largely left to feed its own population.
The probability for the future of food production is that there will be one of the many combinations of the best
and worst in which both are represented.
Whatever that future, one prospect is as certain as anything can be: the developed countries hold virtually all the
cards and will continue to do so. They now have about 2.5 times as much cropland per inhabitant as the
developing countries and in 2050 could have about 4.5 times as much. They have better research facilities and


4
technologies to improve production methods than most developing countries. For example the ability to monitor
within-field spatial variability in soil, crops and environmental factors makes it possible to target inputs to field
crops according to very locally determined requirements. Such technological innovations in agriculture could
benefit developed and developing countries equally, but the cost of setting up such a system makes it more
likely to be used in developed regions.
In the last resort the consumption of animal products in developed countries can be reduced, thereby releasing
more land for the cultivation of crops destined for direct human consumption.
Grain provides at least twice as much food energy when it is consumed directly by humans as when it is fed to
livestock that produce meat and dairy products. Typically, there are big disparities in estimates of the saving of
food by giving up meat and dairy products. A more likely solution to the impending world food crisis would be
for some multi-national organisation on the lines of NATO or the EU to buy cheap grain and other agricultural
commodities and send them to appropriate developing countries, where it would be obtainable on rations at
artificially low prices, a potential problem as already noted, however, for the farmers in such developing
countries.
Views differ widely as to the prospects for the expansion of agricultural production in the next century. In an
ideal world, in which agriculture is managed everywhere at the most efficient possible level and the food needs
of every citizen are catered for, the quantity of food produced could be changed in two ways: either the area of
agricultural land could be extended, or yields could be increased, or both, Since the total land surface of the
world is likely to change only fractionally in the next half century, any increase in the area used for agricultural
purposes would be at the expense of other uses. For example, a 'Good' future for agriculture could mean a 'Bad'
future for the world's forests. The increase in yields needed could not be achieved exclusively by organic
farming and would therefore require, among other changes, a greater application of chemical fertilisers, with
implications for the environment.
To feed some 10 billion people in 2050 will require fuller use to be made of water and land resources, together
with improved means of production and management. The main victims will be much of the present agricultural
population of developing countries, the excessive number of whom needs to be reduced by anything from half to
a tenth, depending on which parts of the world are considered.
2. Livestock production
In 1950 the total number of livestock units in the world was 923 million, giving 366 units per 1,000 people. A
peak of about 430 units per 1,000 people was reached around 1955, after which the number declined to 320 in
1990, when there were about 1,700 livestock units in the world. It means the increase of population was more
intensive compared to the increase of the livestock units.
In order to maintain the 1990 per capita level of livestock units the number of animals should increase steeply in
the future. This projection would put great pressure on arable production and at the same time would require a
considerable improvement in the yield per unit of area of the permanent pasture of the world.
3. Fishing
The fishing industry has been a major source of food and other products for some countries of the world, and is
widely assumed to have a great future potential. Compared with the land, however, the sea offers only a very
limited pasture and in most areas sustains a low density of fish. Between 1930 and 1991 the world fish catch
has increased about ten times form 10 million tonnes to 100 million tonnes as fishing vessels and equipment
have grown in sophistication. During that time the per capita catch increased from 4.6 kilograms to 18.6. If the
trend of 1950-1990 continuous, there is real chance that some sea and ocean areas will be overfished. The
oceans of Antarctica are probably the most extensive area still capable of yielding larger catches.
It is of course a matter of speculation as to how the production of cereals and other crops. Livestock farming and
fishing will change in the next decades. The expected 86% increase in the population of the world until 2050
will no doubt somehow be fed, but with difficulty, possibly with commitment from developed countries with a
comparatively generous endowment of cultivated land and good pasture to supply areas of the present
developing countries unable to cope with population growth. In 2050 the expected per capita arable land gap
between the developed and developing countries of 0.46 and 0.1 hectares per person needs a real solution.


5
Questions:
1. What kind of parameters can have impact on the growth of the worlds population in the future?
2. Evaluate the worlds water resources and their potential change!
3.Describe the trend; the arable land area will change in the different regions of the world!
4. What kind of strategy would be necessary in order to assure enough food for the worlds population in the
future?
4. The animal and its food
Food is material which, after ingestion by animals, is capable of being digested, absorbed and utilised. In a more
general sense we use the term 'food' to describe edible material. Grass and hay, for example, are described as
foods, but not all their components are digestible. Where the term 'food' is used in the general sense, as in this
book, then those components capable of being utilised by animals are described as nutrients.
The animals associated with man cover the spectrum from herbivores, the plant eaters (ruminants, horses and
small animals such as rabbits and guinea pigs); omnivores, which eat all types of foods (pigs and poultry); to
carnivores, which eat chiefly meat (dogs and cats). Under the control of man these major classes of animal still
pertain, but the range of foods that animals are now offered is far greater than they might normally consume in
the wild (for example, ruminants are given plant by-products of various human food industries and some dog
foods contain appreciable amounts of cereals). Nevertheless, plant and plant products form the major source of
nutrients in animal nutrition.
The diet of farm animals in particular consists of plants and plant products, although some foods of animal
origin such as fishmeal and milk are used in limited amounts. Animals depend upon plants for their existence
and consequently a study of animal nutrition must necessarily begin with the plant itself.
Plants are able to synthesise complex materials from simple substances such as carbon dioxide from the air, and
water and inorganic elements from the soil. By means of photosynthesis, energy from sunlight is trapped and
used in these synthetic processes. The greater part of the energy, however, is stored as chemical energy within
the plant itself and it is this energy that is used by the animal for the maintenance of life and synthesis of its own
body tissues. Plants and animals contain similar types of chemical substances, and we can group these into
classes according to constitution, properties and function. The main components are shown in Fig. 1.
Figure 2.3. Fig. 1. The main components of foods, plants and animals

5. Water
The water content of the animal body varies with age. The newborn animal contains from 750 to 800 g/kg water
but this falls to about 500 g/kg in the mature fat animal. It is vital to the life of the organism that the water
content of the body be maintained: an animal will die more rapidly if deprived of water than if deprived of food.
Water functions in the body as a solvent in which nutrients are transported about the body and in which waste
products are excreted. Many of the chemical reactions brought about by enzymes take place in solution and
involve hydrolysis. Because of the high specific heat of water, large changes in heat production can take place
within the animal with very little alteration in body temperature. Water also has a high latent heat of
evaporation, and its evaporation from the lungs and skin gives it a further role in the regulation of body
temperature.
The animal obtains its water from three sources: drinking water, water present in its food and metabolic water,
this last being formed during metabolism by the oxidation of hydrogen-containing organic nutrients. The water
content of foods is very variable and can range from as little as 60 g/kg in concentrates to over 900 g/kg in some
root crops. Because of this great variation in water content, the composition of foods is often expressed on a dry


6
matter basis, which allows a more valid comparison of nutrient content. This is illustrated in Table 3. which lists
a few examples of plant and animal products.
Figure 2.4. Table 3. Composition of some plant and animal products (g/kg)

The water content of growing plants is related to the stage of growth, being greater in younger plants than in
older plants. In temperate climates the acquisition of drinking water is not usually a problem and animals are
provided with a continuous supply. There is no evidence that under normal conditions an excess of drinking
water is harmful, and animals normally drink what they require.
The dry matter (DM) of foods is conveniently divided into organic and' inorganic material, although in living
organisms there is no such sharp distinction. Many organic compounds contain mineral elements as structural
components. Proteins, for example, contain sulphur, and many lipids and carbohydrates contain phosphorus.
It can be seen from Table 3. that the main component of the DM of pasture grass is digestible carbohydrate and
fibre, and this is true of all plants and many seeds. The oilseeds, such as sunflower are exceptional in containing
large amounts of protein and lipid material. In contrast the carbohydrate content of the animal body is very low.
One of the main reasons for the difference between plants and animals is that, whereas the cell walls of plants
consist of carbohydrate material, mainly cellulose, the walls of animal cells are composed almost entirely of
lipid and protein. Furthermore, plants store energy largely in the form of carbohydrates such as starch and
fructans, whereas an animal's main energy store is in the form of lipid.
The lipid content of the animal body is variable and is related to age, the older animal containing a much greater
proportion than the young animal. The lipid content of living plants is relatively low.
In both plants and animals, proteins are the major nitrogen-containing compounds. In plants, in which most of
the protein is present as enzymes, the concentration is high in the young growing plant and falls as the plant
matures. In animals, muscle, skin, hair, feathers, wool and nails consist mainly of protein.
Like proteins, nucleic acids are also nitrogen-containing compounds and they play a basic role in the synthesis
of proteins in all living organisms. They also carry the genetic information of the living cell.
The organic acids that occur in plants and animals include citric, malic, fumaric, succinic and pyruvic acids.
Although these are normally present in small quantities, they nevertheless play an important role as
intermediates in the general metabolism of the cell. Other organic acids occur as fermentation products in the
rumen, or in silage, and these include acetic, propionic, butyric and lactic acids.
Vitamins are present in plants and animals in minute amounts, and many of them are important as components
of enzyme systems. An important difference between plants and animals is that, whereas the former can
synthesise all the vitamins they require for metabolism, animals cannot, or have very limited powers of
synthesis, and are dependent upon an external supply.


7
The inorganic matter contains all those elements present in plants and animals other than carbon, hydrogen,
oxygen and nitrogen. Calcium and phosphorus are the major inorganic components of animals, whereas
potassium and silicon are the main inorganic elements in plants.

8
Chapter 3. Analysis of foods
Originally the most extensive information about the composition of foods was based on a system of analysis
described as the proximate analysis of foods, which was devised over 100 years ago by two German scientists,
Henneberg and Stohmann. Recently, new analytical techniques have been introduced and the information about
food composition is rapidly expanding. However, the system of proximate analysis still forms the basis for the
statutory declaration of the composition of foods in Europe.
1. Proximate analysis of foods
This system of analysis divides the food into six fractions: moisture, ash, crude protein, ether extract, crude fibre
and nitrogen-free extractives. The moisture content is determined as the loss in weight that results from drying a
known weight of food to constant weight at 100 oC. This method is satisfactory for most foods, but with a few,
such as silage, significant losses of volatile material may take place.
The ash content is determined by ignition of a known weight of the food at 550 oC until all carbon has been
removed. The residue is the ash and is taken to represent the inorganic constituents of the food. The ash may,
however, contain material of organic origin such as sulphur and phosphorus from proteins, and some loss of
volatile material in the form of sodium, chloride, potassium, phosphorus and sulphur will take place during
ignition. The ash content is thus not truly representative of the inorganic material in the food either qualitatively
or quantitatively.
The crude protein (CP) content is calculated from the nitrogen content of the food, determined by a modification
of a technique originally devised by Kjeldahl over 100 years ago. In this method the food is digested with
sulphuric acid, which converts to ammonia all nitrogen present except that in the form of nitrate and nitrite. This
ammonia is liberated by adding sodium hydroxide to the digest, distilled off and collected in standard acid, the
quantity so collected being determined by titration or by an automated colorimetric method. It is assumed that
the nitrogen is derived from protein containing 16 per cent nitrogen, and by multiplying the nitrogen figure by
6.25 (i.e. 100/16) an approximate protein value is obtained. This is not 'true protein' since the method determines
nitrogen from sources other than protein, such as free amino acids, amines and nucleic acids, and the fraction is
therefore designated crude protein.
The ether extract (EE) fraction is determined by subjecting the food to a continuous extraction with petroleum
ether for a defined period. The residue, after evaporation of the solvent, is the ether extract. As well as lipids it
contains organic acids, alcohol and pigments. In the current official method, the extraction with ether is
preceded by hydrolysis of the sample with sulphuric acid and the resultant residue is the acid ether extract.
The carbohydrate of the food is contained in two fractions, the crude fibre (CF) and the nitrogen-free extractives
(NFE). The former is determined by subjecting the residual food from ether extraction to successive treatments
with boiling acid and alkali of defined concentration; the organic residue is the crude fibre.
When the sum of the amounts of moisture, ash, crude protein, ether extract and crude fibre (expressed in g/kg) is
subtracted from 1000, the difference is designated the nitrogen-free extractives. The crude fibre fraction
contains cellulose, lignin and hemicelluloses, but not necessarily the whole amounts of these that are present in
the food: a variable proportion, depending upon the species and stage of growth of the plant material, is
contained in the nitrogen-free extractives. The nitrogen-free extractives fraction is a heterogeneous mixture of
all those components not determined in the other fractions. It includes sugars, fructans, starch, pectins, organic
acids and pigments, in addition to those components mentioned above.
2. Modern analytical methods
In recent years the proximate analysis procedure has been severely criticised by many nutritionists as being
archaic and imprecise, and in the majority of laboratories it has been partially replaced by other analytical
procedures. Most criticism has been focused on the crude fibre, ash and nitrogen-free extractives fractions. The
newer methods have been developed to characterise foods in terms of the methods used to express nutrient
requirements. In this way, an attempt is made to use the analytical techniques to quantify the potential supply of
nutrients from the food. For example, for ruminants, analytical methods are being developed that describe the
supply of nutrients for the rumen microbes and the host digestive enzyme system.

Analysis of foods

9
3. Starch and sugars
Inadequacies in the nitrogen-free extractives fraction have been addressed by the development of methods to
quantify the non-structural carbohydrates, which are mainly starches and sugars. Sugars can be determined
colorimetrically after combination with a reagent such as anthrone. Starch is determined by dilute acid
hydrolysis of the sample followed by polarimetric determination of the released sugars. This gives a figure for
total sugars (i.e. those originating from the hydrolysed starch plus the simple sugars in the food). Sugars per se
are determined by extracting the sample with ethanol, acidifying the filtrate and taking a second polarimeter
reading. The starch content is calculated from the difference between the two readings multiplied by a known
factor for the starch source.
4. Fibre
Alternative procedures for fibre have been developed by Van Soest. The neutral-detergent fibre (NDF), which is
the residue after extraction with boiling neutral solutions of sodium lauryl sulphate and ethylenediamine-
tetraacetic acid (EDTA), consists mainly of lignin, cellulose and hemicellulose and can be regarded as a
measure of the plant cell wall material. The analytical method for determining NDF was originally devised for
forages, but can also be used for starch-containing foods provided an amylase treatment is included in the
procedure. By analogy with the nitrogen-free extractives fraction discussed above, the term non-structural
carbohydrate (NSC) is sometimes used for the fraction obtained by subtracting the sum of the amounts (g/kg)
of CP, EE, ash and NDF from 1000.
The acid-detergent fibre (ADF) is the residue after refluxing with 0.5 M sulphuric acid and cetyltrimethyl-
ammonium bromide, and represents the crude lignin and cellulose fractions of plant material but also includes
silica. The determination of ADF is particularly useful for forages as there is a good statistical correlation
between it and the extent to which the food is digested.
The acid-detergent lignin determination involves the preparation of acid-detergent fibre as the preparatory step.
The ADF is treated with 72 per cent sulphuric acid, which dissolves cellulose. Ashing the residue determines
crude lignin, including cutin.
In monogastric, and particularly human, nutrition the term dietary fibre is often used. This is defined as lignin
plus those polysaccharides that cannot be digested by monogastric endogenous enzymes. By virtue of its
definition, dietary fibre is difficult to determine in the laboratory and the alternative term non-starch
polysaccharides (NSP) has been adopted. The NSP in most foods, along with lignin, are considered to represent
the major components of cell walls. The major constituents of NSP are rhamnose, arabinose, xylose, glucose,
galactose, mannose and glucuronic and galacturonic acids. Cellulose is the major source of glucose, and
hemicellulose provides xylose, mannans and galactose. The degradation of pectins releases arabinose, galactose
and uronic acids. A comparison of the dietary fibre contents for a range of food types is given in Table 4.
Figure 3.1. Table 4. The fibre components of some common foods (g/kg D.M.)

In recent years attention has focused on the importance of both water-soluble and insoluble NSP in the human
diet. Water-soluble NSP is known to lower serum cholesterol, and insoluble NSP increases faecal bulk and
speeds up the rate of colonic transit. This last effect is thought to be beneficial in preventing a number of
diseases including cancer of the bowel.
The NSP of foods may be degraded in the gut of pigs by microbial fermentation, yielding volatile fatty acids,
which are absorbed and contribute to the energy supply. A further benefit relates to the volatile fatty acid,
butyric acid, which is reported to be an important source of energy for the growth of cells in the epithelium of
the colon. Thus the presence of this acid will promote development of the cells and enhance absorption. The
extent of degradation depends on the conformation of the polymers and their structural association with non-

Analysis of foods

10
carbohydrate components, such as lignin. In addition the physical properties of the NSP, such as water-holding
capacity and ion exchange properties, can influence the extent of fermentation. The gel-forming NSPs, such as
-glucan, reduce the absorption of other nutrients from the small intestine and depress digestibility and
adversely affect faecal consistency in pigs and poultry. On a positive note, the water-holding properties lead to
beneficial effects on the behaviour of pregnant sows by increasing time spent eating and resting due to increased
gut fill and reducing inappropriate behaviour, such as bar chewing.
5. Minerals
A simple ash determination provides very little information about the exact mineral make-up of the food and,
when this is required, analytical techniques involving spectroscopy are generally used. In atomic absorption
spectroscopy, an acid solution of the sample is heated in a flame and the vaporised atoms absorb energy, which
brings about transitions from the ground state to higher energy levels. The source of energy for this transition is
a cathode lamp, containing the element to be determined, which emits radiation at a characteristic wavelength.
The radiation absorbed by the atoms in the flame is proportional to the concentration of the element in the food
sample.
Flame emission spectroscopy measures the radiation from solutions of the sample heated in air/acetylene or
oxygen/acetylene flames. Each element emits radiation at specific wavelengths and there are published tables of
flame emission spectra. Atomic absorption and flame emission spectrometry are being replaced by inductively
coupled plasma emission spectroscopy, as this has a greater sensitivity for the relatively inert elements and can
be used to determine several elements simultaneously or sequentially.
Just as with other nutrients, a measure of the concentration of the element alone is not sufficient to describe its
usefulness to the animal. Attempts have been made to assess the availability of minerals using chemical
methods, such as solubility in water or dilute acids, but these have had little success. At present animal
experiments are the only reliable way to measure mineral availability.
6. Amino acids, fatty acids and sugars
Knowledge of the crude protein content of a food is not a sufficient measure of its usefulness for non-ruminants.
The amino acid composition of the protein is required in order to assess how a food can meet the essential
amino acid requirements. Similarly, the total ether extract content does not give sufficient information on this
fraction since it is important to know its fatty acid composition. In non-ruminants, this has large effects on the
composition of body fat and, if soft fat is to be avoided, the level of unsaturated fatty acids in the diet must be
controlled. In ruminants, a high proportion of unsaturates will depress fibre digestion in the rumen. When
detailed information on the amino acid composition of protein, the fatty acid composition of fat or the individual
sugars in NSP is required, then techniques involving chromatographic separation can be used. In gas-liquid
chromatography, the stationary phase is a liquid held in a porous solid, usually a resin, and the mobile phase is a
gas. Volatile substances partition between the liquid and the vapour and can be effectively isolated. This form
of chromatography is, however, usually a slow process and in order to speed up the separation procedure, high-
performance liquid chromatography has been developed. In this technique, pressure is used to force a solution,
containing the compounds to be separated, rapidly through the resin held in a strong metal column. In addition
to speeding up the process, high resolution is also obtained. Gas-liquid chromatography and high-performance
liquid chromatography can also be used for the determination of certain vitamins (e.g. A, E, B6, K), but the
measurement of available vitamins requires biological methods.
An example of the application of high-performance liquid chromatography is seen with food proteins, which are
hydrolysed with acid and the released amino acids are then determined using either:
1. Ion exchange chromatography - by which the amino acids arc separated on the column, then mixed with a
derivatisation agent, which reacts to give a complex that is detected by a spectrophotometer or fluorimeter.
2. Reverse phase chromatography - in which the amino acids react with the reagent to form fluorescent or
ultraviolet-absorbing derivatives, which are then separated using a more polar mobile phase (e.g. acetate buffer
with a gradient of acetonitrile) and a less polar stationary phase (e.g. octadecyl-bonded silica). The availability
of amino acids to the animal can be estimated by chemical methods. For example, for lysine there are
colorimetric methods which depend on the formation of compounds between lysine and dyes.

Analysis of foods

11
7. Measurement of protein in foods for ruminants
The new methods of expressing the protein requirements of ruminants require more information than just the
crude protein (nitrogen) content of the food. In the metabolisable protein system, used in Hungary the
unavailable nitrogen is measured as acid detergent insoluble nitrogen (ADIN). For the determination the acid
detergent extractions of Van Soest, described above, is used. It is the only parameter among protein fractions,
can be determined by a laboratory method. Information on the rate of degradation in the rumen of the available
nitrogen is also required and this can be estimated by biological methods.
8. Spectroscopy
More recently, in some laboratories, procedures combining statistical regression techniques with near-infrared
reflectance spectroscopy (NIRS) have been introduced. Energy in the wavelength range from 1100 to 2500 nm
is directed on to a cell containing the dried milled sample, and the diffuse reflected energy is measured. This is
related to the chemical nature of the sample, since the bonds -CH, -OH, -NH and -SH absorb energy at specific
wavelengths. A spectrum of reflected energy is recorded and, by use of a computer, the spectral data obtained
from a calibration set of samples are related by multiple linear regression to their known composition
determined by traditional methods. These relationships are then used to relate the reflectance readings obtained
for individual foods to their composition. This technique is now used routinely to determine a range of food
characteristics, including those that are the resultant of a number of nutrient concentrations such as digestibility,
metabolisable energy and nitrogen degradability in the rumen.
Questions:
1. What are the major differences in the nutrient content of plants and animals?
2. What are the main nutrient categories of the wendee analytical system?
3. What are the main faults of the wendee system?
4. What kind of other analytical categories and procedures do you know?

12
Chapter 4. The significance of
antinutritional factors
Many feedstuffs that are commonly used in preparing diets for animals contain antinutritional factors (ANFs).
These factors interfere with the utilisation of dietary nutrients in a variety of ways, including reducing protein
digestibility, binding to various nutrients or damaging the gut wall and thereby reducing digestive efficiency.
The main ANFs that interfere with nutrient digestion and absorption are lectins, protease inhibitors, tannins,
antigenic proteins, phytic acid, glucosinolates and gossypol. For some ANFs, lectins and tannins in particular,
their proper characterisation and analysis are the main limitations towards a better understanding of their effects
on animals. The effects of lectins, trypsin inhibitors and tannins on apparent and true amino acid digestibilities
have been documented. The application of heat substantially reduces the activity of several ANFs, and in
particular lectins and protease inhibitors. An effective means to improve the availability of phosphorus bound to
phytic acid is the use of exogenous phytases. For tannins and glucosinolates, no practical means for inactivation
are yet available. More information is needed before we can routinely quantify levels of ANFs in feedstuffs and
relate ANF contents to effects on true nutrient digestibilities and gut endogenous nutrient losses.
Many feedstuffs that are commonly fed animals contain factors that interfere with the utilisation of dietary
nutrients. These factors, which cause depressions in growth and feed efficiency and/or affect animal health, can
be defined as antinutritional factors (ANFs). In plants and seeds these ANFs primarily act as biopesticides,
protecting them against moulds, bacteria and birds.
In this chapter the following aspects of ANFs will be discussed: classification and occurrence of ANFs in
feedstuffs; methods of chemical analysis; the effects of ANFs on animals; and some of the means to inactivate
or reduce negative effects of ANFs. Emphasis will be placed on the effects of ANFs on the digestion of energy
yielding nutrients and amino acids. The latter effects may allow us to relate ANF contents to true nutrient
digestibilities and endogenous nutrient losses, even though our knowledge of quantitative effects of ANFs on
protein and energy utilisation is far from complete. According to the above definition, dietary fibre (the non-
starch polysaccharides, NSPs), can also be classified as antinutritional.
ANFs can be classified in various ways. The following classification, based on their effects on the nutritive
value of feedstuffs and on biological responses in animals, can be suggested:
- Factors that have a depressive effect on protein digestion and utilisation (trypsin and chymotrypsin inhibitors,
lectins or haemagglutinins, polyphenolic compounds, NSP-s and saponins).
- Factors which have a negative effect on the digestion of carbohydrates (amylase inhibitors, polyphenolic
compounds, NSP-s, flatulence factors).
- Factors which have a negative effect on the digestion and utilisation of minerals (glucosinolates, oxalic acid,
phytic acid, gossypol).
- Factors which inactivate vitamins or cause an increase in the animal's vitamin requirements (anti-vitamins ).
- Factors that stimulate the immune system (antigenic proteins).
The type and content of ANFs in different types of feedstuffs vary considerably. Moreover, many feedstuffs
contain several ANFs, and the amounts of ANF can vary considerably between batches of the same feedstuff.
The latter variation can be attributed to the plant's growing conditions as well as to genetics; different varieties
can have different levels of ANF. In legume seeds (soya beans, peas and beans) protease inhibitors and lectins
are the most important ANFs. However, some varieties of cereal grains, rye and triticale in particular, may also
contain moderate levels of trypsin inhibitors. Tannins are mainly present in the coloured-flowering varieties of
vicia faba beans, peas, rapeseed (canola), sorghum and some varieties of barley. Glucosinolates and sinapins are
important in rapeseed; alkaloids in lupins, and gossypol in cottonseed.
1. Lectins
Lectins, or haemagglutinins, are proteins that are generally present in the form of glycoproteins. They vary
considerably in their molecular weight and chemical structure and are characterised by an ability to bind to

The significance of antinutritional
factors

13
specific sugars. Glycoproteins in the gut wall contain sugars to which lectins have affinity and, as a result,
binding of lectins to epithelial cells occurs. This leads to growth depressions. A prerequisite for the
antinutritional properties of lectins is resistance to proteolysis. A variety of lectins exist, both within and
between different types of plant seed, which have differing effects on the animal. In general, the lectins in
common beans are highly toxic, while lectins in peas and faba beans appear to be the least toxic. Furthermore,
different animals may respond to the same lectins in different ways; piglets appear to be more sensitive than rats
and chickens. Haemagglutination of red blood cells is most commonly used to measure lectin activity. This is
based on the ability of lectins to bind to sugars on the surface of the red blood cells. However, this assay appears
not to be very specific and it is inaccurate in predicting the effects of lectins on animals. More specific assays
that have been developed recently include ELISA and functional lectin immunoassay (FLIA). The latter is based
on the ability of lectins to bind to a specified carbohydrate matrix or a gut wall brush border membrane
preparation. Due to considerable variations in chemical structure between lectins from different legume seeds, a
specific assay (ELISA or FLIA) is required for each legume seed. When purified soya bean lectins are included
in pig diets they increase endogenous gut nitrogen losses as measured at the terminal ileum. (Table 5.). These
endogenous losses, probably arising from a loss of mucus, do not appear to increase in proportion to the lectin
content of the diet. Lectins appear to have minor effects on true protein digestibilities but reduce the digestibility
of other nutrients as indicated by the greater flow of dry matter at the distal ileum in pigs given lectins (Table 5.
).
Figure 4.1. Table 5. Effects of feeding soya bean lectins on flows of dry matter and
nitrogen at the distal ileum in growing pigs (Schulze et al., 1995)

An additional effect of lectins is that they stimulate the proliferation of bacteria in the intestinal lumen. The
exact reason for this is not clear, although it may be related to increased nutrient availability to the bacteria and
an increase in epithelial cell turnover, which may then increase the number of potential binding sites for bacteria
on epithelial cells.
As lectins are quite unstable to heat, heat processing appears to be an effective means of inactivating lectins.
2. Protease inhibitors
There are various categories of plant protease inhibitors. The main inhibitors in legume seeds and cereals are the
trypsin and chymotrypsin inhibitors. These are peptides that form stable inactive complexes with some of the
pancreatic enzymes. As a result the activities of trypsin and chymotrypsin are reduced. Inactivation of these
enzymes in the gut induces the endocrine cells in the mucosa to release more cholecystokinin (CCK) which
stimulates the pancreas to produce more digestive enzymes. In particular, feeding protease inhibitors to rats and
chickens results in hypertrophy of the pancreas. Furthermore, other feed constituents, such as tannins, can
inhibit trypsin activity. Therefore, care should be taken in the interpretation of trypsin inhibitor activity (TIA)
values. In a survey of the literature, TIA in raw peas was about 12% of that in raw soya beans, while that in
common beans was about 38% of the activity in soya beans. Low protease inhibitor activity occurs in cereal
grains such as rye, triticale, wheat, barley and oats. It should be noted that there are important differences in
trypsin inhibitor content between different varieties of the same seed. For example, soya bean varieties that are
low in specific anti-trypsins have been developed. Trypsin inhibitor units (TIU/mg) vary between 1.69 and
11.24 in different varieties of raw peas. Trypsin inhibitors substantially increased the ileal flow of both
endogenous and exogenous nitrogen, as determined using the 15N-isotope dilution technique. Even though only
a limited number of levels of TIA were evaluated. Schulze (1994) suggested that trypsin inhibitors affect
endogenous nitrogen losses and true nitrogen digestibility, in a linear manner. Similar observations were made
by Barth et al. (1993) feeding a synthetic diet to miniature pigs. The addition of graded levels of Kunitz trypsin-
inhibitor reduced apparent ileal protein digestibility from 74.5% (no added trypsin inhibitor) to 56% (3000 mg
added TIA per meal). In contrast to Schulze et al. (1995), Barth et al. (1993) concluded that the reduction in
apparent protein digestibility could be attributed to increases in endogenous protein losses rather than to
reductions in true ileal protein digestibilities. This apparent discrepancy may be related to the different varieties
of soya beans that were evaluated or to differences in the techniques used to quantify endogenous protein losses.
Negative relationships between TIA in peas and apparent faecal protein digestibilities have been observed in
pigs (Leterme et al. 1989), rats (Birk 1989), and poultry (Carre & Conan 1989). However in the latter study, less

factors

14
than 25% of the variability in protein digestibility in six pea samples could be explained by the anti-trypsin
content in the samples. Obviously, additional factors influence apparent protein digestibility in peas. Based on a
review of the literature, van Leeuwen et al. (1993) estimated that endogenous ileal protein losses increase by
nine grams for each unit of extra trypsin inhibitor activity in peas. The impact of this increase in endogenous
losses on observed protein digestibilities will vary with the protein level in the feed ingredient.
Trypsin inhibitors are less susceptible to heat than lectins. However, and as shown in Table 6., a moderate heat
treatment is an effective means to inactivate soya bean TIA. However, heat treatment needs to be conducted
under closely controlled conditions to avoid reducing the availability of amino acids, particularly lysine.
Figure 4.2. Table 6. Effcet of alternative heat treatments on lectin and trypsin inhibitor
activity in phaseolus beans (van der Poel, 1999)

Feeding previously germinated feedstuffs known to contain high levels of ANFs reduces endogenous gut protein
loss (Table 7.). A significant reduction in ileal crude protein loss at the terminal ileum occurs as a result of
germinatio.
Figure 4.3. Table 7. Contents of antinutritional factors and crude protein digestibility
values of raw and germinated beans, measured with growing pigs

3. Alpha-amylase inhibitor
The alpha-amylase inhibitor is thought to be responsible for the impaired digestion of starch in red kidney
beans. However, this ANF appears to be of minor importance, as its addition in a purified form to a mixed diet
does not impair starch digestion.
4. Tannins
Tannins are polyphenolic compounds with a range of molecular weights and variable chemical complexity.
They are capable of precipitating alkaloids and gelatine as well as other proteins from aqueous solutions.
Although tannins are chemically not well defined, they are usually divided into two subgroups: hydrolysable
and condensed tannins. Hydrolysable tannins have a central carbohydrate core the hydroxyl groups of which are
esterified to various phenolic carboxylic acids. This group of tannins is easily hydrolysed to give glucose or a
polyhydroxy alcohol and the various phenolic acids.

factors

15
A variety of assays are available to quantify tannin contents in feedstuffs . However, most of these assays are
not very specific or quantify only one component of the tannins. The most commonly used approach is the
modified vanillin method in which tannin content is expressed as catechin equivalents (Price et al. 1978). One of
the main limitations towards a better understanding of the effects of tannins on animals is a lack of suitable
analyses and characterisation.
The amount of tannins in sorghum varies considerably from undetectable levels, to levels close to 11%. In
selected samples of cottonseed meal tannin content ranged from 8 to 15 g of condensed extractable tannins per
kg.
The way in which tannins affect animal performance is not exactly clear. Tannins form complexes with proteins
and carbohydrates in the feeds, and with digestive enzymes. As a result nutrient digestibility is depressed. Other
effects of tannins include reduced feed intake, increased damage to the gut wall, toxicity of absorbed tannins and
reduced absorption of some minerals. These effects can largely be attributed to condensed tannins. However,
absorption of hydrolysable tannins or their degradation products appear to have direct effects on the liver and
possibly other organs. The fact that no clear relationships between tannin levels in sorghum and depressions in
growth performance in birds have been established, appears to support the notion that we are unable to properly
quantify the (toxic) tannin content in feedstuffs. Furthermore, some of the depressions in performance due to the
intake of tannins can be overcome by increasing dietary protein levels. This also complicates the interpretation
of animal performance studies where effects of dietary tannins are evaluated. In terms of nutrient digestibilities,
significant differences were found in ileal and faecal nutrient digestibilities in piglets fed different cultivars of
faba beans (Table 8.). This depression in apparent protein digestibility can be attributed equally to increases in
endogenous N losses and to increases in the excretion of dietary protein at the distal ileum. However, when
these same varieties were fed to broiler chickens between 5 and 26 days of age and at similar inclusion levels,
no differences in growth performance were observed. This again illustrates that piglets are more sensitive to
tannins than broiler chickens.
Figure 4.4. Table 8. Apparent ileal and faecal digestibilities of crude protein and
nitrogen free extract in different cultivars of faba beans to piglets, included at 300g/kg
(Jansmann, 1993)

Animals and humans that consume high tannin diets develop a physiological means to counteract the adverse
effects of tannins. This occurs through the production of a family of proline-rich proteins in the salivary gland.
These proteins bind tannins in a highly specific manner, thereby reducing their toxicity. However, pigs and birds
are unable to completely eliminate the toxic effects of dietary tannins. Alternative means to detoxify tannins
include the addition of chemicals (such as polyethylene glycol) that have high affinity for tannins in the diet,
soaking of feedstuffs in water or alkaline solutions, anaerobic fermentation or germination of sorghum.
However, none of these methods has been proven to be cost-effective.
5. Flatulence factors
Flatulence is a digestive disorder that is characterised by an abnormal amount of gas formation in the
gastrointestinal tract following consumption of oligosaccharides that are not digested by mammals and birds.
The major flatulence producing factors are raffinose, verbascose and stachyose all of which belong to the
raffinose family of oligosaccharides (RFO). The fermentation of RFOs in the hindgut results in the production
of carbon dioxide, hydrogen, and methane as well as volatile fatty acids. At high concentrations these products
result in flatulence, diarrhoea, nausea, clamp, and general discomfort to the animal.

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The problem of flatulence has been studied more in relation to human nutrition. Various ways of overcoming
the development of flatulence have been studied. Cooking, soaking, fermentation, dehulling and enzyme
supplementation procedures have been shown to decrease the flatulence properties of cowpeas and the same is
likely true for other pulse crops and legumes. On a commercial scale, it will be economically attractive to use
enzyme preparations as opposed to soaking, fermentation and dehulling because the latter procedures will
require redrying of the ingredients before they are used in diet preparation.
6. Antigenic proteins
Antigenic proteins are macromolecular proteins or glycoproteins capable of inducing a humoral immune
response when fed to animals. A humoral immune response occurs when specific polyclonal antibodies are
produced and secreted into body fluids such as the blood. Antigenic proteins are known to cause gut wall
damage and immunological reactions in the gut linked with disorders in gut function in piglets and veal calves.
A major concern with immune responses is the development of chronic hypersensitivity to antigenic
compounds. In particular, hypersensitivity occurs in calves fed milk replacers containing soya bean products.
This can result in increased endogenous gut protein secretions and changes in gastro-intestinal motility and
morphology as well as gut permeability. Fortunately, most laboratory animals and pigs can develop a tolerance
to antigenic proteins that are present in the diet.
Antigens do not appear to be very sensitive to heat. Chemical and enzymatic treatments appear to be more
effective ways to reduce antigenicity in soya bean products. For this reason, soya concentrates that are included
in calf milk replacers are ethanol/water-extracted products.
7. Phytic acid
Phytic acid is the acid form of the anion phytate (myo-inositol hexa phosphate). Phytic acid can not be
hydrolysed by enzymes secreted into the gut by mammals and birds. Phosphorus present in phytic acid has a law
bio-availability. Moreover, phytate can form complexes with a variety of minerals, including calcium, copper,
cobalt, icon, magnesium, manganese, selenium and zinc, thus reducing the availability of these nutrients. Phytic
acid can also form complexes with basic residues of proteins and therefore it may interfere with the activity of
endogenous enzymes and digestibilities of nutrients other than minerals.
The content of phytic acid in various feedstuffs and its effect on the availability of phosphorus has been
documented. Furthermore, the use of exogenous phytases to enhance phosphorus digestibility is now common
practice in countries where the contribution of animal agriculture to environmental pollution is a concern. The
effects of phytases on amino acid and energy digestibilities as well as endogenous nutrient losses have not been
adequately investigated. However, there are indications that phytases enhance apparent ileal amino acid and
faecal energy digestibilities in pigs and poultry. Recent studies have shown that phytase supplementation greatly
increases the body retention of nitrogen, calcium and phosphorus. In the study by Mroz et al. (1994), in which
growing-finishing pigs were fed a corn-tapioca-soya bean meal diet, the addition of 800 phytase units per kg of
diet, led to a reduced excretion of nitrogen, calcium and phosphorus by 5.5%, 2.2% and 1.9%, respectively.
8. Vicine and convicine
Vicine and convicine are glycosides that are primarily found in faba beans. These compounds are hydrolysed by
the intestinal microflora to divicine and isourarnil These degradation products cause haemolytic anaemia, in
man. In birds they result in a decrease in egg weight and size, weaker egg shells, an increased number of blood
spots in the egg, and a decrease in fertility and hatchability of eggs. In pigs they have been related to reduced
reproductive performance. Vicine and convicine, and their degradation products appear to have no direct effect
on nutrient digestion and metabolism. The levels of vicine and convicine vary considerably between varieties of
faba beans; the most effective means to reduce the levels of these ANFs in faba beans is the selection of
varieties with law contents.
9. Saponins
Saponins are steroid or triterpenoid glycosides that are present in many feedstuffs. They have a bitter taste, can
form foams in aqueous solutions and haemolyse red blood cells. They are known to depress growth performance
in both poultry and swine. Their antinutritional properties seem related to their ability to form complexes with

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sterols, in particular those in membranes of animal cells. This appears to result in increased permeability of the
intestinal mucosa. Poultry, compared to other monogastrics, are more sensitive to saponins. Significant saponin
levels are present in alfalfa meal with minor levels in other legumes such as soya beans, rapeseed and various
varieties of peas. In general saponins are of minor concern in monogastric animals because they are present at
only law levels in common feedstuffs.
10. Glucosinolates
Glucosinolates are present in all cruciferous seeds and plants. Over a hundred glucosinolates have been
identified. Of these, less than ten are of concern to animal nutritionists, in particular those present in rapeseeds
and rapeseed meal. Glucosinolates can be categorised into two main groups: the aliphatic types and indole types.
Their quantities in feedstuffs can be determined by gas chromatography. These ANFs are hydrolysed by the
enzyme myrosinase present in cruciferous seeds and plants, although bacteria that are present in the gastro-
intestinal tract also contribute to the hydrolysis of glucosinolates. Hydrolysis of glucosinolates yields glucose,
various goitrogenic compounds (thiocyanates, isothiocyanates and oxazolidinethione), and nitriles. The
thiocyanate ions in particular, inhibit the uptake of iodine by the thyroid gland for the production of
triiodothyronine (T3) and thyroxine (T4) leading to lower plasma levels of these compounds. Nitriles are known
to cause lesions in the liver and kidney of rats, poultry and pigs. The severity of the effects of glucosinolates or
their degradation products in animals is influenced by a number of factors including species, age and the growth
status of the animal. Glucosinolates and their degradation products do not appear to have a direct effect on
nutrient digestion and absorption.
The effects of glucosinolates can be overcome by the use of supplemental copper sulphate or iodine in the diet.
However, the most promising means to reduce effects of glucosinolates is reducing their content in feedstuffs
through plant breeding. Although development of double- or triple-low cultivars of rapeseed has eliminated the
palatability problems, concerns remain regarding the inclusion levels of rapeseed or rapeseed meal in animal
diets. This is indicative of the fact that the role of glucosinolates as ANFs with respect to their effects on the
thyroid, metabolism and growth, is poorly understood. Consequently, the use of rapeseed or rapeseed meal in
diets for young animals should be avoided and there is need for the development of plant varieties with even
lower levels of glucosinolates.
11. Oxalic acid
Oxalic acid is a primary toxic constituent in many plants. It binds with divalent cations such as calcium and
magnesium thus decreasing their utilisation. Excessive intakes of oxalic acid can cause diarrhoea, gastroenteritis
and renal damage.
12. Gossypol
Gossypol is a yellow, phenolic pigment found in the glands of cottonseed from the genus Gossypium. The
analysis of free gossypol appears to have a high repeatability. The gossypol content of cottonseed varies
considerably between different varieties and growing conditions, with levels ranging from 0.002 to 6.64% being
reported. High temperatures and pressures during processing favour the formation of stable bonds between
gossypol and protein which may be responsible for reduced availability of amino acids, and lysine in particular,
in cottonseed meal. Furthermore, gossypol can cause death if dietary levels exceed 0.015%, the highest level of
free gossypol considered safe for pigs. Other effects of high dietary levels of free gossypol include respiratory
distress, abdominal distension and infertility in male pigs. Mortality rates due to gossypol poisoning can be as
high as 80% in piglets. Some of the adverse effects of gossypol can be prevented by providing supplemental
iron in the diet.
13. Alkaloids
Alkaloids are compounds that contain nitrogen in a heterocyclic ring, are generally basic and often have a bitter
taste. The word alkaloid simply means 'alkali-like'. Alkaloids are present in many plants whereby they are
thought to serve as a chemical defence against herbivory. Lupins in particular, contain high levels of alkaloids,
specifically quinolizidine alkaloids. In some instances soya beans and linseed may be contaminated with Datura
stramonium. The latter seed contains the alkaloids hyoscyamine and scopolamine.

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Alkaloids are oxidized in the liver resulting in metabolites, such as dehydrosparteine, which are responsible for
the observed toxicity. The level of toxicity is influenced by the structure of the alkaloids. There is a high degree
of variation in the ability of different animal species to deal with these compounds. The toxic effects of alkaloids
and their metabolites are primarily mediated through the nervous system, although they are also stimulate
copper uptake by liver cells thus leading to copper toxicity. The negative effect of lupins on feed intake appears
to be mediated through the neural system as well. Pigs appear more sensitive to alkaloids than poultry. New
varieties of lupins have become available that contain low alkaloid levels as well as low levels of other ANFs
such as trypsin inhibitors and lectins.
14. Sinapins
Sinapins are present in rapeseed meal at varying levels. These compounds are converted to trimethylamine in
the hindgut and contribute to taint (fishy odour) in brown shelled eggs. Layers producing white shell eggs and
mammals appear to be able to degrade trimethylamine. Sinapins do not interfere with nutrient digestion or
metabolism.
15. Non-starch polysaccharides
Over 100 monosaccharides are found in nature but only about nine of these are the predominant building blocks
of NSP. These include the pentoses (arabinose and xylose), hexoses (glucose, mannose and galactose), the 6-
deoxyhexoses (rhamnose and fucose) and hexauronic acids (galacturonic acid and glucuronic acid). The number
of potential polysaccharide structures is enormous. The complexity is further increased by covalent bonding to
non-carbohydrate compounds, such as methyl and acetyl groups, phenolic acids, proteins and lignin, and non-
covalent bonding within and between polysaccharides. This diversity in chemical structure, both within and
between dietary sources, is reflected in physical structure and the physiological activities of various NSP
fractions.
The main NSP structures, commonly found in feed ingredients of plant origin, are listed in Figure 2.
Figure 4.5. Figure 2. Polysaccharide structures commonly found in feed ingredients of
plant origin (de Lange, 2000)

Cellulose is present in most plants as the structural polysaccharide in plant cell walls. In cereals, the main matrix
polysaccharides in cell walls are beta-glucans in barley and a variety of arabinoxylans in wheat, rye, sorghum,
corn and triticale. The NSP structures present in legumes are more complex than those in cereals.
In addition to the NSPs listed in Figure 2. some feedstuffs contain substantial amounts of oligosaccharides that
cannot be degraded by enzymes produced by mammals and birds. The primary oligosaccharides are -
galactosides (raffinose, stachyose and verbascose), which contain three to nine monomers of galactose and
glucose linked via -1,6 bonds. Substantial quantities are present in soya bean meal (about 6%), peas (5%), and
rapeseed meal (3%).
It can be argued that starch and sugars that are not enzymatically digested, have similar properties to NSPs.
Three main fractions of resistant starch (RS) can be identified: physically inaccessible starch (RS1), uncooked
native starch granules (RSz) and retrograded starch (RS3). It is clear that the three fractions of resistant starch
can be manipulated by processing of feeds and adding NSP degrading enzymes.

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16. Mechanisms by which dietary NSPs affect animal
performance
NSPs have varying effects on animals when included in the diet. They may affect the production and activity of
digestive enzymes, intestinal morphology (including cell proliferation), the microbial population in various
segments of the gut, and secretion of certain hormones, including insulin, glucagon.
NSPs, unlike starch (-glucans) and sucrose (glucose and fructose connected via a (-1,4 bond), cannot be
hydrolysed by enzymes produced by mammals or birds themselves. For the digestion and utilisation of energy
containing material in NSPs, animals are dependent on microbial fermentation in the intestinal lumen. This
fermentation results in the production of short-chain fatty acids, which may be absorbed and used as a source of
energy. The digestibilities of NSPs are lower than those of starch and sugars. Furthermore, the efficiency of
utilisation of energy derived from digested NSP is about 30% lower than that of starch and sugars. This has
direct implications for the supply of effective energy to the animal. In this context, resistant starch (including
sugars) that is not digested enzvmatically, but that is fermented, should also be quantified.
In various studies, the negative effects of increased dietary NSP levels on the digestibility and rate of absorption
of nutrients from starch, protein and fats have been demonstrated (1995; Bakker 1996; Smits 1996). Several
mechanisms are involved in reducing the rate and extent of apparent nutrient digestion and absorption. First, the
endogenous secretions and losses of enzymes and mucus and the sloughing of mucosal cells are likely increased
when intake of NSP is increased. Furthermore, endogenous secretions, such as bile acids, can be bound,
particularly by viscous or gelling and lignified NSP, thus reducing the extent of recycling. These effects result in
reductions in apparent nutrient digestion. Second, viscous NSP in particular, will interfere with digesta
movement and the mixing of digestive enzymes and nutrients in the intestinal lumen. Combined with increased
mucus production, NSPs can also increase the resistance of the unstirred water layer at the intestinal surface.
Third, NSPs from cell walls can physically hinder the access of digestive enzymes to nutrients that are enclosed
inside cell walls. Fourth, soluble NSPs in particular may stimulate microbial growth and increase the amount of
microbial protein and fat at the terminal ileum or in faeces. Selected NSPs may also stimulate the growth of
toxin producing microbes, which may affect gut health directly and digestive function indirectly. Fifth, feeding
animals NSPs may alter intestinal morphology and the capacity of the gut to absorb nutrients.
In poultry in particular and more specifically in broiler chickens, many of these negative effects on nutrient
digestibilities have been related to the viscosity of digesta when feeding increasing levels of NSP. An increased
viscosity of digesta is induced primarily by soluble viscous NSPs, such as arabinoxylans, beta-glucans and
pectins. For example, Annison (1991) found a clear inverse relationship between in vivo soluble NSP extract
and AME in Australian wheat samples fed to broiler chickens. It should, however, be noted that similar
relationships could not be established for UK wheat (McNab 1992; Austin et al. 1998). This apparent
discrepancy may be explained by the difference in structure of the arabinoxylan, the main soluble NSP in wheat,
in different wheat varieties.
The effect of NSPs on the viscosity of digesta is influenced by factors such as feed processing, the presence of
endogenous hydrolytic enzymes in the plant material, exogenous enzymes, as well as microbial fermentation
and enzymatic digestion in the intestine.
The effects of soluble, non-viscous NSPs and oligosaccharides (e.g. -galactosides) on poultry are not clearly
established. In some instances no effect has been observed (Trevino et al. 1990), while in other situations
negative effects (Leske et al. 1995) on broiler chick performance have been reported.
In pigs, and other simple stomached animals, the relationship between solubility and/or viscosity and the
antinutritional properties of NSPs are not as clear, or not as clearly established, as reported for poultry (Graham
& Aman 1991). For example, Larsen et al. (1994) did not observe differences in apparent ileal amino acid and
nitrogen digestibility in rats fed varying levels of a synthetic viscous NSP. Furthermore, in a range of wheat
samples there was no strong relationship between the soluble NSP content and digestible energy (DE) values
(Zijlstra et al. 1999). This likely relates to differences in digestive physiology between monogastric mammals
and poultry. In particular, the longer retention time of digesta in pigs increases the opportunity for (viscous)
NSPs to be fermented and nutrients to be absorbed.
This implies that the effect of soluble and/or viscous NSPs on the ileal digestibiliry of nutrients in pigs may be
small, or less important in pigs than in poultry. More data are required to relate dietary soluble and/or viscous
NSP contents to ileal and faecal nutrient digestibilities for pigs and other monogastric mammals. It appears that

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more conventional measures of NSPs, such as crude fibre (CF) and neutral detergent fibre (NDF), provide for a
reasonable means to predict the effect of NSPs on nutrient and energy digestibilities in various classes of pig
feed ingredients. The latter suggests that in the pig the effects of NSPs on nutrient digestibility are due largely to
physical enclosure of dietary nutrients and possibly to effects on gut health.
Monogastric animals attempt to increase feed intake when given diets that contain increasing levels of NSP.
This is probably an attempt to maintain the rate of available energy intake.
A reduction in the amount of fermentable substrates entering the hindgut of pigs or broiler chicks may improve
aspects of gut health. The effects of NSPs on gut microflora are partly confounded with effects on digesta
viscosity and water holding capacity. As a result, it is difficult to establish whether effects on viscosity and
water holding capacity, or effects on gut microflora, are the main means by which NSPs reduce nutrient
digestibility. It should be noted that specific NSPs (-glucans) or oligosaccharides may be used to enhance gut
health in young pigs. These 'pre-biotic' properties may be due to direct effects on intestinal microbes.
Alternatively, they may be targeted towards specific binding sites on the intestinal mucosa and, as such, reduce
the attachment of (harmful) microbes to the intestinal surface.
In summary, the digestible NSP content of a feedstuff should be determined when evaluating its available
energy content. Furthermore, diet NSP contents have considerable effects on digestibility of other dietary
nutrients, voluntary feed intake, metabolism of nutrients after absorption, and gut health. These effects can be
attributed to the effects of NSPs on gut microflora, viscosity and water-holding capacity of the digesta, and to
the 'mechanical' properties of the NSP. There are likely to be threshold values below which NSP levels do not
affect animal performance. In fact, at low inclusion levels, selected NSPs or oligosaccharides may stimulate
digestive function and improve gut health.
Questions:
1. What are the main groups of anti-nutritive substances in feedstuffs?
2. Which feedstuffs are especially concerned?
3. What are main effects of anti-nutritive materials on the production of farm animals?
4. What kind of procedures can be used for inactivating the anti-nutritive materials of feedstuffs?

21
Chapter 5. Feed additives
Feed additives are materials that are administered to the animals to enhance the effectiveness of nutrients and
exert their effects in the gut or on the gut wall cells. There are clear theoretical reasons why these additives
should be effective but, given the dynamic nature of the gut physiology, it is often very difficult to demonstrate
the effects in practice. For example, the effects of added organic acids within the gut lumen are difficult to
quantify, even when re-entrant cannulae are employed.
1. Antibiotics
Antibiotics are chemical compounds which, when given in small amounts, halt the growth of bacteria. They are
produced by other microorganisms, e.g. fungi, and are also synthesised in the laboratory. They are used at
therapeutic levels, by injection or in food or water, to treat diseases caused by bacteria. In addition, sub-
therapeutic levels of antibiotics are added to the food to enhance the rate of growth. The various groups of
antibiotics act in different ways to reduce the numbers of specific bacteria in the gut, and thereby increase the
efficiency of nutrient utilisation. This is brought about by:
- reduction or elimination of the activity of pathogenic bacteria which may cause subclinical infection, thus
allowing the host to achieve production levels closer to the potential;
- elimination of those bacteria which produce toxins that reduce the growth of the host animal;
- stimulation of the growth of microorganisms that synthesise unidentified nutrients;
- reduction of the growth of microorganisms that compete with the host animal (the fermentation of nutrients by
bacteria is a wasteful process compared with direct absorption);
- increased absorptive capacity of the small intestine through a decrease in the thickness of the intestinal wall.
These effects may be coupled with a reduced turnover of mucosal cells and reduced mucous secretion. The gut
accounts for a large proportion of the energy and protein required maintaining an animal and any reduction in
the mass of the gut and cell turnover will release nutrients for other purposes such as growth.
Antibiotic growth promoters have been used mainly in pig and poultry foods, typically at levels of 20-40 mg/kg,
which give improvements of 4-16 per cent in growth rate and 2-7 per cent in efficiency of food conversion. The
response is greatest in young animals and those consuming diets containing vegetable protein rather than animal
protein. Mortality in young pigs is also reduced. The effect is less in healthy herds and flocks. Young pre-
ruminant calves also respond to growth-promoting antibiotics in the same manner as non-ruminants. Since
ruminants depend primarily on bacteria in the rumen for their nutrient supply, the use of antibiotics in ruminant
diets might be considered to be disadvantageous. However, certain antibiotics of the ionophore type (e.g.
monensin sodium) have been found to be effective, particularly with low-roughage/high-concentrate diets, in
controlling the proportion of undesirable bacteria in the rumen without disturbing the overall fermentation.
There are small changes in the fermentation, however, such as reduced methane production and increased
propionic acid proportions, which improve productivity. Monensin sodium at 20-30 mg/kg of food improves the
efficiency of food conversion by increasing the rate of gain with the same food intake or by maintaining the rate
of gain at a lower food intake. Amino acid degradation is decreased and the additional propionic acid spares
amino acids for gluconeogenesis.
The widespread use of antibiotics coupled with the ability of resistant bacterial strains to evolve over a short
period of time and transfer resistance to other strains has resulted in populations of bacteria that are resistant to
many antibiotics. This has limited the effectiveness of antibiotic treatment against disease and, whereas at one
time a bacterial infection could be treated with a range of antibiotics, now some bacteria are sensitive to a single
antibiotic only. There is concern that if such bacteria gained resistance to this antibiotic then the disease would
be untreatable. For this reason the use of antibiotics as growth promoters has been curtailed by legislation in
recent years. In the EU in 2000, only four antibiotics were permitted for use as growth promoters:
fIavophospholipol, avilamycin, salinomycin sodium and monensin sodium.
Another material which enhances the growth of pigs, and is thought to act via the antibacterial effects described
above, is copper . When added to their food, as copper sulphate, at levels far in excess of the requirement for
normal functions (e.g. 200 mg Cu/kg compared with 5 mg Cu/kg) pigs grow faster. Concern over the

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22
accumulation of copper in the environment and the potential toxic effect of applying manure of high copper
content to pasture grazed by sheep, has resulted in legislation in the EU to control the amount of copper that
may be added to pig foods. The food of young pigs up to 16 weeks of age may contain 175 mg Cu/kg but this
must be reduced to 100 mg/kg thereafter, and from six months of age the maximum level is 35 mg/kg.
2. Modes of action antibiotics
Antibiotics halt the growth of bacteria by interfering with their cellular metabolism.
There are four groups:
1. Antibiotics which interfere with the synthesis of the material forming the bacterial cell wall and cause the cell
to burst. These are high molecular weight (> 1200) compounds that act on Gram-positive bacteria. They are
poorly absorbed by the host and thus are non-toxic, leave no detectable residues and have no withdrawal period
(i.e. a period of time during which the compound must be removed from the food before the animal is
slaughtered). Avoparcin and flavomycin are examples of this type of antibiotic.
2. Inhibitors of bacterial protein synthesis are also primarily active against Gram-positive bacteria and have a
medium molecular weight (>500). Although they are absorbed to a greater extent than the higher molecular
weight compounds, they do not have a withdrawal period. Examples of this type of antibiotic include tylosin and
virginiamycin.
3. Inhibitors of bacterial DNA synthesis can have a broad spectrum of activity, have a law molecular weight
(about 250) and require withdrawal periods. Nitrofurans and quinoxaline-N-oxides fall into this category of
antibiotics.
4. Ionophore antibiotics interfere with the electrolyte balance (Na/K) of the bacterial cell by transporting
potassium into the cell, which then has to use energy to pump it out. Eventually the ion pump fails to operate
efficiently and potassium accumulates inside the cell. Water enters by osmosis and the cell ruptures. Monensin
sodium is an example of this type of antibiotic.
3. Probiotics
In contrast to the use of antibiotics as nutritional modifiers, which destroy bacteria, the inclusion of probiotics in
foods is designed to encourage certain strains of bacteria in the gut at the expense of less desirable ones. A
probiotic is defined as a live microbial food supplement that beneficially affects the host animal by improving
the intestinal microbial balance. Although the digestive tract of all animals is sterile at birth, contact with the
mother and the environment leads to the establishment of a varied microflora. The beneficial microorganisms
produce enzymes which complement the digestive ability of the host and their presence provides a barrier
against invading pathogens. Digestive upsets are common at times of stress (e.g. weaning), and feeding with
desirable bacteria, such as Lactobacilli, in these situations is preferable to using antibiotics, which destroy the
desirable bacteria as well as the harmful species.
It has been suggested that the desirable bacteria exert their effects in a number of ways:
- Adhesion to the digestive tract wall to prevent colonisation by pathogenic microorganisms. Detrimental
bacteria, such as E. coli, need to become attached to the gut wall to exert their harmful effects. Attachment is
achieved by means of hair-like structures, called fimbriae, on the bacterial surface. The fimbriae are made up of
proteins, called lectins, which recognise and selectively combine with specific oligosaccharide receptor sites on
the gut wall. Lactobacilli successfully compete for these attachment sites (Fig. 3. )
- Neutralisation of enterotoxins produced by pathogenic bacteria which cause fluid less. There is some evidence
that live probiotic bacteria can neutralise these toxins but the active substance has not been identified.
- Bactericidal activity: Lactobacilli ferment lactose to lactic acid, thereby reducing the pH to a level that harmful
bacteria cannot tolerate. Hydrogen peroxide is also produced, which inhibits the growth of Gram-negative
bacteria. It has also been reported that lactic acid producing bacteria of the Streptococcus and Lactobacillus
species produce antibiotics.

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- Prevention of amine synthesis: coliform bacteria decarboxylate amino acids to produce amines, which irritate
the gut, are toxic and are concurrent with the incidence of diarrhoea. If desirable bacteria prevent the coliforms
proliferating then amine production will also be prevented.
- Enhanced immune competence: oral inoculation of young pigs with Lactobacilli has elevated serum protein
and white blood cell counts. This may aid the development of the immune system by stimulation of the
production of antibodies and increased phagocytic activity.
Figure 5.1. Fig. 3. Competitive exclusion of pathogen bacteria due to attachment of non-
pathogens

Other postulated effects include beneficial interaction with bile salts, increased digestive enzyme production and
more efficient absorption of nutrients, and greater vitamin production. In a review of the response of pigs of
various ages to the administration of probiotics it was concluded that they were effective for young pigs, in
which the digestive tract is still developing after weaning. However, they were less effective for growing and
finishing pigs, which already have a balanced population of microorganisms. To be effective the desirable
microorganism should not be harmful to the host animal, should be resistant to bile and acid, should colonise the
gut efficiently, should inhibit pathogenic activity, and should be viable and stable under manufacturing and
storage conditions.
In monogastric animals, strains of Lactobacilli, Bacillus subtilis and Streptococci have been used as probiotics.
In ruminant animals, the application of yeast (Saccharomyces cerevisiae) in the form of live culture, or dead
cells with culture extracts, has proved successful in beneficially modifying rumen fermentation.
Yeast cultures stimulate forage intake by increasing the rate of digestion of fibre in the rumen in the first 24
hours after its consumption. Overall digestibility is not affected. It is likely that this improvement in early
digestion and intake is brought about by alterations in the numbers and species of microorganisms in the rumen.
The precise means by which the effect is achieved have not yet been confirmed but there are a number of
probable mechanisms.
It is thought that metabolites of dead and live yeast cells (B vitamins, branched chain fatty acids, amino acids
and peptides) stimulate the growth of the bacterial species Megasphaera elsdenii. This utilises the lactic acid
produced from the rapid fermentation of starch and sugars associated with high-concentrate diets. Live yeasts
ferment sugars derived from the degradation of starch, thus competing with the lactic acid producing bacteria,
and thereby stabilise rumen pH and reduce the risk of acidosis. Live yeast cultures also scavenge oxygen in the
rumen, helping to maintain anaerobic conditions and favouring the growth of cellulolytic bacteria. The increase
in forage intake results in improved live weight gain, milk yield and milk fat content. Improved fibre digestion
has also been reported in horses when yeast cultures have been given.
4. Prebiotics

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Oligosaccharides (2-20 monosaccharide units) have been claimed as beneficial nutritional modifiers for
monogastric farm animals. They fall into the group of materials also known as prebiotics, which are defined as
compounds, other than dietary nutrients, that modify the balance of the microfloral population by promoting the
growth of beneficial bacteria and thereby provide a healthier intestinal environment. Oligosaccharides occur
naturally in foods: soya bean meal, rapeseed meal and legumes contain -galactooligosaccharides (GOS);
cereals contain fructooligosaccharides (FOS); milk products have trans-galactooligosaccharides (TOS); yeast
cell walls contain mannanoligosaccharides (MOS). They are also produced commercially.
It has been suggested that these compounds achieve their beneficial effects in the gut in two ways. First,
although they are not easily digested by the host digestive enzymes, compounds such as FOS can be fermented
by the favourable bacteria (e.g. Bifidobacteria and Lactobacilli), giving them a competitive advantage. This
shifts the microbial population towards such microorganisms and away from the harmful species.
Second, the gut microbial population may be altered by the oligosaccharide interfering with the attachment of
harmful bacteria to the gut wall. As a means of cell recognition all cell types have a unique configuration of
carbohydrate-containing compounds (glycoproteins and glycolipids) on their surface. As described above in the
section on probiotics, pathogenic bacterial cells have surface compounds called lectins which recognise these
carbohydrates and by which they attach to the gut cells. Once attached the bacteria are able to multiply and
produce their harmful effects. Species such as Salmonella and E. coli have a mannose-specific lectin which
binds to mannose residues on the gut mucosal surface. By introducing mannose-containing compounds (MOS)
into the diet the binding by pathogenic bacteria is disrupted and instead they bind to the oligosaccharide and are
carried out of the gut with the passage of the digesta (Fig 4.). Yeasts have mannans in the cell wall structure and
form the basis of some commercial products that are claimed to act in this way. Indeed the presence of such
yeast fragments has been said to be the reason why yeast products are beneficial.
Figure 5.2. Fig. 4. Effect of oligosaccharides on the excretion of harmful bacteria

The efficacy of products containing oligosaccharides is currently the subject of active experimentation. There
can be no guarantee that an oligosaccharide will favour the growth of beneficial species in a complex microflora
such as that found in the pig's intestine. Experiments have shown that piglets given an oral challenge of E. coli
responded to GOS with a reduced pH of ileal digesta and reduced population of coliforms. Supplements of FOS
and TOS have reduced the numbers of aerobic bacteria in the gut of weaned piglets and there are reports of a
reduced incidence of diarrhoea. Under farm conditions, improvements in gain and food conversion efficiency of
the order of 4-6 per cent have been recorded. In other experiments reduced digesta pH has been reported but
without a detectable change in the composition of the microflora, microbial metabolites or production responses.
These conflicting results may have arisen because the diet already contained some oligosaccharides or because
experimental conditions tend to be less stressful than those on farms.
5. Enzymes
Not all compounds in foods are broken down by the enzymes so some potential nutrients are unavailable. The
chemical structure of plant cells (fibre), the limited time available for activity in certain parts of the gut and the
presence of antinutritive compounds in some foods hinder the release of nutrients. Early experiments with
supplemental enzymes to overcome these limitations yielded variable responses, mainly because they were
impure materials. As a result of advances in biotechnology, more effective enzyme preparations can now be
produced in large quantities and relatively inexpensively. Therefore, supplementation of the diet as a means of
improving nutritive value is becoming commonplace.
The enzymes used as food additives act in a number of ways. First, they can improve the availability of plant
storage polysaccharides (e.g. starch), oils and proteins, which are protected from digestive enzymes by the
impermeable cell-wall structures. Thus cellulases can be used to break down cellulose, which is not degraded by
endogenous mammalian enzymes. Even in ruminant animals, in which the rumen microflora possess cellulase,
fibrolytic enzymes have been used but responses have been variable owing to the complex nature of the rumen
fermentation system and other factors such as lignin encrustation of cellulose. Responses in pigs and poultry of
the order of 5-10 per cent improvement in live weight gain and 10 per cent improvement in food conversion

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efficiency have been recorded. Supplementation of a wheat by-product diet with cellulase increased the ileal
digestibility of non-starch polysaccharides from 0.192 to 0.359 and crude protein from 0.65 to 0.71.
Second, enzymes are employed to destroy materials that interfere with the digestion, absorption and utilisation
of nutrients. The major application in this area has been to break down carbohydrate fractions in cereal cell
walls, the -glucans and arabinoxylans, which are resistant to attack by digestive enzymes. These materials
hinder the digestion and absorption of other nutrients by forming a viscous gum in the digestive tract. In pigs, it
has been suggested that the increased viscosity brought about by -glucans in the stomach contents may affect
the natural sieving of particles. Thus large particles are suspended in the viscous digesta and pass through to the
duodenum instead of falling to the bottom of the stomach, resulting in less efficient digestion. It is also possible
that the increase in viscosity disturbs peristalsis and pancreatic secretion. In poultry, the viscous nature of the
digesta results in poor performance and sticky droppings, which present a problem in the management of litter
waste. Supplements of -glucanase to barley-based diets improved the rate of gain and food conversion
efficiency of broilers to values similar to those for wheat- or maize-based diets (Table 9.). The -glucanase also
reduced litter problems and improved the metabolisable energy of the diet.
Figure 5.3. Table 9. Effect of -glucanase addition to barley on the production
parameters of 3 week old broiler chicks

Supplementation of pig foods improves ileal and whole tract digestibility of nutrients. In breaking down the -
glucans, the enzymes reduce the viscosity of the digesta thus allowing better movement of endogenous enzymes
through the mass and more efficient digestion and absorption of nutrients. It is still not clear whether viscosity
per se is responsible for the effects or if it is an indicator of conditions pertaining in the gut lumen which cause
the problems. The response differs between pigs and poultry because of differences in digestive physiology
between the two species.
Phytase is in this category of enzyme. It releases the orthophosphate groups from phytic acid, which is only
partially broken down by non-ruminants and is the major form of phosphorus in cereal grains and oilseeds.
These results in a greater availability of phosphorus to the animal and the amount of inorganic phosphorus
added to the diet can be reduced with beneficial effects on the environment through reduced phosphorus
excretion.
In the third area of enzyme application, the aim is to supplement the enzyme complement of young animals, in
which the rate of endogenous enzyme production may be limiting. Early weaned pigs have limited amylase,
protease and lipase activity and enhancement of the extent of digestion of nutrients would improve performance
and reduce the incidence of the diarrhoea that results from undigested nutrients reaching the hind gut and being
fermented by bacteria.
Finally, enzymes can increase the efficiency of utilisation of nutrients, in monogastric animals, by releasing
them for uptake from the small intestine rather than allowing fermentation in the hind gut, which results in
products of lower value to the animal (e.g. volatile fatty acids).
In order to be effective when incorporated into the animal's diet, enzymes must survive storage at ambient
temperature, the manufacturing process (heating and pelleting) and wide fluctuations in pH in the gut, be
resistant to intestinal proteases and have specific activity on feed components in the upper digestive tract. The
enzyme should be selected on the basis of its target substrate. Commercial enzymes are derived from a wide
range of sources, particularly fungi such as Trichoderma longibrachiatum, Aspergillus niger and Humicola
insolens.
6. Enzyme action in pigs and poultry

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Enzymes have found major commercial application in pig and poultry diets and are extensively used for the
latter. Differences in efficiency have been reported between the species, the effect generally being greater in
poultry; this is possibly due to the differences in gut physiology between the two species. There is a greater
reduction in the viscosity of digesta in the small intestine by enzymes in poultry, probably because of the lower
water content than in pigs. The longer retention time in the stomach and acidic nature of stomach contents in the
pig (around pH 3) is more destructive to enzymes than the milder conditions in the crop (pH 4-5). New
commercial enzymes have been selected for their pH stability. Finally, pigs have a longer small intestine than
poultry and the resultant increased retention time allows more time for the endogenous enzymes to act. Thus
there is less opportunity to detect the effects of supplemental enzymes in pigs.
7. Organic acids
Efficient digestion of food in young pigs depends on the secretion of sufficient acid in the stomach. Acid
additives have been used mainly in the diets of early weaned pigs. Here their use relates to the digestive
physiology of the young pig and the replacement of sow's milk with solid food at weaning. The suckled piglet
obtains nutrients on a little-and-often basis, sucking about once every hour. The milk clots in the stomach, and
the limited secretion of hydrochloric acid is augmented by lactic acid produced from the fermentation of lactose
by Lactobacilli. This ensures that the pH falls to a level that favours efficient protein hydrolysis and suppresses
harmful bacteria. Thereafter there is a steady release of nutrients from the clot to the small intestine. The solid
food presented at weaning is less digestible than milk, is consumed in larger, less frequent meals and has a
higher buffering capacity. The limited hydrochloric acid production is not sufficient to ensure that the pH falls
to a level which prevents the growth of pathogenic bacteria. In addition the nutrients may overload the immature
digestive and absorptive capacity of the small intestine and their fermentation in the hind gut results in
diarrhoea.
In order to overcome these problems acids have been added to weaning diets to augment gastric HCI, giving a
rapid fall in pH with beneficial effects on protein digestion and the gut microflora. Although inorganic acids
have been used to supply hydrogen ions, and hence reduce pH, organic acids have additional desirable
properties. In their salt form they are odourless and easy to handle; they lower the pH and acid binding capacity
of the food; the anion has an antimicrobial effect on moulds and bacteria in the food, thereby improving food
quality. The anion has been reported to act as a complexing agent for cations, such as Ca++ and Mg++, thus
improving their retention. It is also suggested that the presence of short chain fatty acids in the small intestine
reduces the damage to the villi of the gut wall which is associated with weaning. Finally, organic acids, once
absorbed by the pig, can be used as substrates in intermediary metabolism, principally as energy sources.
It has proved difficult to obtain experimental evidence for all of these purported effects of organic acids. In an
extensive review Partenen and Mroz (1999) examined the evidence for their modes of action. Only a few studies
have demonstrated a reduction in gastric pH, but the methodology of sampling is difficult given diurnal
variations, differing proportions of food and secretions, and timing of sampling. A study is required with
permanent fixed pH electrodes within the stomach. There is evidence of a positive effect on the ileal
digestibility of amino acids and the retention of nutrients in growing pigs. The magnitude of the effect depends
on the acid used and its concentration, the age of the pig and the composition of the diet. With regard to
minerals, the absorption of calcium and phosphorus has been improved but variable results have been obtained
for retention of these minerals. Addition of organic acids has had beneficial effects on microbial counts and
species in some studies but there has been little work on the effects on gut morphology to test the theory of an
improvement of the structure of the villi. Finally, in some studies, the addition of formic, fumaric and citric
acids to the diet has increased the activity of enzymes associated with intermediary metabolism, indicating that
the acids do contribute to nutrient metabolism.
Although the exact modes of action of the acids are not clear, their addition to pig diets has proved beneficial in
terms of nutrient digestibility, growth and food conversion efficiency (Table 10. ). This is so, particularly for
newly weaned pigs, when coupled with good feeding management procedures. Formic and propionic acids are
more effective than fumaric or citric acids at the same rate of inclusion because the former have a lower
molecular weight. Suggested levels of inclusion of acid (kg/tonne diet) are: formic 6 to 8; propionic 8 to 10;
fumaric 12 to 15; citric 20 to 25, but recommendations vary.
Figure 5.4. Table 10. Effect of graded levels of fumaric acid on the growth rate and food
conversion efficiency of young pigs. Easter R.A. (1988): Acidification of diets for pigs,
In: Haresign, W. and Cole D.J.A. Recent Advances in Animal Nutrition

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8. Antimicrobial action of organic acids
Organic acids change from the undissociated to the dissociated form, depending on the pH of the environment.
In the undissociated form they can diffuse rapidly through the semipermeable membrane into the cytoplasm of
the microorganism. Once inside, at a pH of around 7, the acid becomes dissociated and acts by suppressing the
cellular enzyme and transport systems. Longer chain length acids and unsaturated acids are the most effective.
9. Modifiers of rumen fermentation
Rumen fermentation processes are usually kept within narrow bounds and, as they form an integrated system,
any changes made have more than one outcome. Thus modification of the processes to one desired target is
difficult. Nevertheless, there are products which are used to modify the fermentation to achieve specific results.
The use of antibiotics and probiotics was discussed above. Other rumen modifiers include buffers, methane
inhibitors and bloat-preventing substances. Buffers are included in ruminant diets to regulate rumen pH to levels
that favour the activity of cellulolytic organisms (pH 6-7). Diets rich in readily fermentable carbohydrate create
acidic conditions and increase lactic acid formation, both of which are detrimental to the cellulolytic bacteria.
This reduces food intake and predisposes the animal to acidosis and laminitis. Secondary problems include
reduced mills fat production, rumenitis, ketosis and liver abscesses. Chemicals such as sodium bicarbonate,
sodium carbonate, calcium carbonate and magnesium oxide buffer the hydrogen ions and increase the dilution
rate of the liquids in the rumen. This increases the efficiency of microbial protein synthesis and, as a result of
the shorter retention time, allows starch and protein to escape to the intestines. The activity of cellulolytic
bacteria and the proportion of acetate in the rumen volatile fatty acids are increased. In the USA it has become
common to add up to 200 g NaHCO3/day, and sometimes MgO in addition, to the rations of cows in early
lactation. It has been suggested that, since bicarbonate buffers are expended once the CO2 is released, it would
be advantageous to use certain clays and bentonites, which contain insoluble and hence undegradable anions and
would allow the buffering action to be carried further down the digestive tract.
Antibiotics such as virginiamycin and monensin have been used to select against bacteria that produce lactic
acid. Certain probiotics encourage the uptake of lactic acid.
The production of methane in the rumen is a wasteful process (up to 10 per cent of the gross energy) and
contributes to the earth's so-called `greenhouse gases'. Halogen compounds, particularly bromide and chloride
homologues of methane, are toxic to the bacteria which produce methane. Suppression of methane production
results in a reduction in the NAD/NADH ratio, which is unfavourable to anaerobic fermentation. However, the
accompanying increase in propionate and butyrate production uses up the excess hydrogen, re-establishing the
NAD/NADH ratio. Long chain polyunsaturated fatty acids also reduce methane production. Some studies have
shown partial adaptation by the rumen microflora to these additives.
Under normal conditions rumen fermentation produces gases which, by the action of ruminal and reticular
contractions, accumulate in the area known as the cardia, from which they are passed up the oesophagus in the
process of eructation. Bloat occurs if the gas becomes trapped and the ani-mal is not able to clear it. Rumen
stasis due to acidosis is one of the causes of free gas bloat but on certain pastures, particularly those rich in
legumes, and on high-concentrate diets the fermentation process in the rumen can result in rumen gases being
trapped within fluid bubbles. Therefore, the gas cannot be eructated as normal and pressure builds up in the
rumen. The condition is known as frothy bloat. In the case of legume bloat the frothing is due to chemicals
associated with the plant. With high concentrate diets the normal function of the rumen is disrupted by the rapid
accumulation of fermentation acids and the release of bacterial mucopolysaccharides during cell lysis, which
result in increased viscosity of the rumen fluid and the formation of the stable foam. In the case of pasture bloat

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there are management practices (e.g. strip grazing or use of grass/clover mixtures) to reduce its incidence but
often these are insufficient. Antibiotics have been used to change the end-products of fermentation in order to
produce fewer froth-forming materials. However, specific anti-foaming agents are more commonly used. These
reduce the surface tension of the rumen fluid, preventing the formation of the stable foam. Vegetable oils,
lecithin, animal fats, mineral oils, detergents and the synthetic polymer poloxalene fall into this category of
agent. The additive alkylarylsulphonate prevents the release of pectin derivatives, which cause frothing. The
main problem with such additives is the maintenance of an adequate concentration in the rumen, since liquids
quickly pass through. Spraying the forage with oil has proved successful. In New Zealand, a product
containing platonic detergents and alcohol ethoxylate is used for pasture bloat but its effectiveness, and that of
other pasture bloat remedies, for feedlot bloat is not known. For the latter type of bloat, the effectiveness of the
ionophore antibiotics has been partially attributed to the lower absolute amount and daily variation of food
intake. The addition of sodium chloride at 40 g/kg diet DM has been suggested as a preventive measure as this
increases the rate of passage of fluids from the rumen, but it also depresses food intake.
Questions:
1. Explain the way how probiotics, prebiotics and symbiotics exert their positive effect!
2. What kinds of enzymes are used in the practice of animal nutrition?
3. What are the main advantages of using crystalline amino acids?
4. What kind of anti-oxidants do you know? Under what circumstances can they have especial importance?

29
Chapter 6. Advances in feed
evaluation for pigs
In commercial pork production there is a move towards the development of more precise and situation specific
feeding programmes for individual pig units. This reflects an increase in the accuracy of feed evaluation through
knowledge of the feed and animal factors that influence nutrient utilisation in pigs, an ability to apply this
knowledge effectively in practice through the use of mathematical animal production models, and the impact of
feeding programmes on profits. In this chapter six topics are addressed that ileal with specific aspects of the
evaluation of pig feed ingredients that have implications for formulating pig diets:
1. There is considerable variation in feeding values between different batches from the same feed ingredient. For
various feed ingredients, methods of estimating feeding values from simple measurements have been developed.
Relationships between available energy contents and proximate analyses that have been established across
different feed ingredients may not be sufficiently sensitive to identify accurately differences in available energy
contents between different batches of the same feed ingredient. Some care should be taken when mathematical
equations are used to predict amino acid contents from protein contents, in particular for cereal grains and
probably also for oil seed meals. This is because relationships between protein and amino acid contents are
likely to differ between different plant varieties.
2. In commercial feed compounding, an increasing variety of processing methods is used. The effect of
mechanical and thermal processing on feeding values can be substantial and will vary with types of feed
ingredients. Furthermore, for the laboratory evaluation of nutritive values of feed ingredients, processing may
differ substantially from that used in feed compounding. Processing should thus be considered carefully when
assigning feeding values to feed ingredients.
3. An important prerequisite in feed formulation is that feeding values of individual feed ingredients are
additive. There is considerable evidence to suggest that apparent ileal digestible amino acid values are not
additive when mixing ingredients with high and low amino acid values. For this reason 'standardised' amino acid
digestibilities should be used in which apparent amino acid digestibilities in feed ingredients are corrected for
minimum gut endogenous amino acid losses. In diets with high contents of fat, non-starch polysaccharides or
synthetic amino acids, the available energy content of the diet may not be accurately predicted based on the
available energy values for the individual feed ingredients.
4. Substantial amounts of microbial fermentation occur in the upper gastrointestinal tract. This is likely to have
implications for the energetic utilisation of starch and may interfere with the interpretation of ileal amino acid
digestibilities.
5. Feed ingredients, in particular those that contain large amounts of non-starch polysaccharides and
antinutritional factors, can increase the nutrient and energy requirements by visceral organs. This will reduce the
efficiency with which absorbed available nutrients are used for muscle or lean tissue growth and reproduction.
6. Finally, there are important effects of body weight and stage of production (growing, gestating, lactating) on
determined nutrient digestibilities. Different nutritive values should be assigned to pig feed ingredients for each
stage of production.
In commercial pork production there is a move to more advanced feeding practices, such as phase and split-sex
feeding, feeding to specific animal performance potentials, feeding to minimise the impact on the environment,
and situation-specific feeding programmes to maximise profitability on individual pig units. Furthermore, due to
economic, environmental and public pressures, a wide range of feed ingredients are included in pig diets. It is
thus becoming increasingly important to characterise accurately the nutritional values of pig feed ingredients
and to predict the pig's response when fed various combinations of feed ingredients. Moreover, a wide range of
feed additives (including flavours, acidifiers, antioxidants, enzymes, growth promoters, prebiotics and
probiotics, immune function modulators, sources of antibodies) is available that may enhance nutrient
availability and the utilisation of absorbed available nutrients by pigs. In a similar manner, alternative methods
of processing feed ingredients and complete feeds should be considered.
During the last decade we have seen some major advances in systems that can be used to assess the available
energy and amino acid contents of pig feed ingredients. For example, net energy and true ileal digestible amino

Advances in feed evaluation for pigs

30
acid contents have been determined for a wide range of ingredients. These systems are increasingly used in the
formulation of commercial pig diets, in particular in parts of the world where a wide range of feed ingredients
and by-products from the food industry are used. Moreover, mathematical animal production models are
increasingly being used to develop cost-effective feeding strategies for individual pig units.
In this chapter selected topics that have implications for the routine formulation of pig feeds will be discussed.
These include: variability in feeding value within the same feed ingredient, effects of processing on the feeding
value of feed ingredients, interactive effects of feed ingredients on nutrient digestion and utilisation, feed
ingredient specific effects on nutrient metabolism in visceral organs and differences in nutrient digestibilities
between different classes of pig.
1. Variability in feeding value within the same feed
ingredient
No two batches of a feed ingredient are the same. Variability in the composition of plant feeds such as cereal
grains and legume seeds arises from differences in variety, soil type, climate, use of fertilisers and other aspects
of husbandry, as well as from post-harvest treatment and processing. Feeds of animal origin, such as fish meal
and meat meals are equally subject to variability related to the species used, the parts used and the conditions
under which they were processed. These variations in composition give rise to substantial differences in
nutritional value.
It is not feasible in practical feed compounding, to carry out a complete evaluation of every batch of an
ingredient. Moreover, in vitro techniques or near infrared reflectance analyses are still often considered too time
consuming, too expensive, or not sufficiently accurate. For many nutrients, values will be taken from tables of
feed composition, either published tables or those compiled in-house. Tables of feed composition give average
values for a given ingredient and their use necessarily introduces some error when applied to most samples. To
deal with this, dietary inclusion levels of ingredients that are known to be variable in feeding value may be
limited. Alternatively, stochastic feed formulation systems may be used in which variability in available nutrient
contents of feed ingredients is explicitly represented and the probability that available nutrient contents in the
complete feed exceed minimum levels is specified. To achieve greater accuracy in predicting the feeding value
of ingredients it is necessary to develop methods for estimating the unknown parameters from those that are
known. To some extent this is already done in tables of feed composition, where, for example, values are
tabulated for hard wheat and soft wheat. Building a database containing details of the provenance of feed
samples can greatly assist in assigning more appropriate nutritive values to a particular batch.
Further improvements in assigning accurate nutritional values can be derived from simple chemical and physical
analyses of the material. Many aspects of composition are interrelated. For example, meat meals with low bone
and fat contents contain more lean meat and automatically have higher concentrations of protein and because
they are made with more lean meat and less connective tissue they have higher concentrations of essential amino
acids and higher protein quality. Such interrelationships amongst the components of a feed can be useful
predictors of nutritional value. For selected ingredients, methods to predict digestible or metabolisable energy
contents from simple measurements have been developed. It should be noted that relationships between
digestible, metabolisable or net energy contents and proximate analyses, that have been established across
different feed ingredients, may not be sufficiently sensitive to accurately identify differences in available energy
contents between different batches from the same feed ingredient.
Positive relationships between protein and amino acid contents have been developed for the main feed
ingredients. However, some care should be taken in the interpretation of these relationships. For example, a
simple negative relationship between the nitrogen concentration and the lysine concentration in the grain protein
has been demonstrated for barley as well as for other cereal grains (Fickler et al. 1995) This arises because
increases in nitrogen concentration increase the endosperm protein relatively more than the aleurone proteins,
which tend to have higher concentrations of lysine and other essential amino acids. However, these associations
are found only when increases in grain nitrogen are brought about by fertiliser usage or other environmental
influences. Differences in nitrogen concentration between varieties are not generally associated with such
systematic alterations in amino acid composition (Fuller et al. 1989), though varieties do vary in amino acid
composition, the most extreme examples being those mutants with increased lysine concentrations such as Riso
barley and Opaque-2 maize.
To improve these general predictions of cereal composition requires a better description of the provenance of
samples, for example, the variety, soil type, climate, use of fertilisers and yield of cereals.


31
2. Effects of processing on the feeding value of feed
ingredients
Feed ingredients are often identified simply by name, regardless of how the material is processed either before
or after its incorporation into a diet. In modern feed compounding, an increasing variety of processing methods
is being used. This includes grinding, flaking, micronisation, extrusion, expansion, pelleting, double pelleting,
and extended conditioning prior to pelleting. While processes like grinding and pelleting can be applied to most
feed ingredients, other more advanced and expensive processing methods, such as extrusion, are likely to be
used for specific feed ingredients that will benefit most from these forms of processing. The inputs of
mechanical and thermal work involved can radically change the physical structure and chemical composition of
the ingredient, leading to changes in its properties, such that it is inappropriate to describe it by the same name
as the original material.
Mechanical energy, especially shear energy, tends to comminute the feed particles, exposing a greater surface
area to enzyme attack. Thermal energy inputs alter both physical structure and chemical composition. Any
thermal processing which raises the temperature above approximately 60C results in the disruption of starch
granule structure and leads to gelatinization. Gelatinization makes the starch more immediately available to the
animal's amylolytic enzymes. Plant cell walls are inherently resistant to attack by the digestive secretions of
animals. The actions of thermal energy on cell walls are more complex, having both positive and negative
effects. On the one hand, the disruption of cell walls would make the cell contents more accessible to the
digestive enzymes of the animal. On the other hand, it increases solubilisation of cell wall polysaccharides
(beta-glucans and arabinoxylans). These components of dietary fibre increase the viscosity of digesta and may
impair the digestion of other dietary constituents, such as protein and fat. These effects are particularly
important in poultry but are also seen in pigs, especially young pigs. These effects will differ with the type of
ingredient and interactive effects between different types of processing may exist. These effects can be at least
partially reversed by the addition of exogenous enzymes.
Against this scenario, in commercial feed manufacturing, feed ingredients are normally evaluated under
standard conditions. Samples for in vivo evaluation are usually ground in some standard way and not further
processed. Samples for in vitro evaluation are commonly ground in a laboratory mill, much finer than
conventional hammer-milling. Thus, the sample evaluated is often processed differently from the sample
actually to be fed to the animal.
Accurate evaluation of feed ingredients, therefore, needs to take account of processing and ascribe a different
value according to the processing conditions. This might include the prediction of viscosity and the resultant
changes in digestibility. Addition of exogenous enzymes could be treated analogously.
3. Interactive effects of feed ingredients on nutrient
digestion and utilisation
An important prerequisite in feed formulation is that the feeding values of individual feed ingredients are
additive. In general, this appears to be the case. However, in some specific situations a lack of additivity has
been demonstrated. This lack of additivity may be attributed to interactive effects of feed ingredients on nutrient
digestion and absorption or on the efficiency of nutrient utilisation after absorption. Diets with extreme levels of
fat and non-starch polysaccharides (NSPs), ileal and faecal digestibilities of some nutrients are not predicted
accurately from those in pure ingredients. In particular, at high levels of NSP intake, the faecal output of crude
protein was increased, while the urinary output of nitrogen was reduced. The latter can be attributed to increased
microbial fermentation in the hindgut and a redirection of nitrogen excretion from urine to faeces. However,
when both soya hulls (a source of easily fermentable NSPs) and fat were added to the diet, this effect was no
longer observed. The effects of high dietary NSP and fat levels may be attributed to effects on the viscosity of
digesta, microbial fermentation and possibly anatomical and metabolic changes in intestinal tissue. It should be
noted that these interactive effects were demonstrated at high fat and NSP levels. Furthermore, the type of fat
may affect these interactions. They are lower when oils, rather than animal fats, are included in the diet. Some of
the effects of exogenous dietary enzymes on nutrient utilisation in pigs should be considered in the same
context. Even though exogenous enzymes are targeted towards specific materials (e.g. phytate-phosphorus,
NSPs), they may also result in changes in ileal and faecal digestibilities of other components (amino acids,
starch) in the diet. For example, exogenous phytase not only increases the digestibility of phytate-phosphorus, it
also slightly increases the apparent ileal digestibility of nutrients other than phosphorus.


32
For amino acids, there may be a lack of additivity of apparent ileal amino acid digestibilities when low protein
feed ingredients are combined with high protein feed ingredients (Table 11.).
Figure 6.1. Table 11. Determined and calculated amino acid digestibilities for selected
amino acids

This can largely be explained by the curvilinear effects of dietary amino acid levels on observed apparent ileal
amino acid digestibilities. This, in turn, can be attributed to the contribution of the minimum gut endogenous
amino acid losses to the total flow of amino acids at the terminal ileum. The correction of apparent ileal amino
acid digestibilities for minimum gut endogenous amino acid losses (that are assumed not to be affected by diet
composition) will eliminate these effects. The resulting 'standardised' amino acid digestibilities are more likely
to be additive than apparent digestibilities. Unfortunately, there is considerable variation between estimates of
minimum gut endogenous amino acid losses. The variation may be attributed to differences in methods,
including analytical procedures used to estimate these losses as well as between-animal variation.
Different estimates are used in various tables in which amino acid digestibility values are summarized (CVB
1998; NRC 1998). Extreme care should thus be taken in combining results from different tables. The most
appropriate means to assess whether these minimum gut endogenous amino acid losses are estimated correctly is
to evaluate the relationship between amino acid levels and 'corrected' amino acid digestibilities across studies
and within ingredients; there should be no effect of dietary amino acid level on 'corrected' amino acid
digestibility. An important point in using these 'corrected' ileal amino acid digestibilities in feed formulation is
that the minimum gut endogenous amino acid losses should be taken into account when considering amino acid
requirements. They should be added to the 'maintenance' amino acid requirements when requirements are
estimated factorially.
The efficiency of utilisation of nutrients after absorption is determined by the nutrient levels in the diet and the
purpose for which these nutrients are used (body 'maintenance' functions, body protein and body lipid
deposition, or milk protein and milk lipid production). For example, when absorbed dietary fatty acids and
amino acids are used to generate high energy phosphate bonds in adenosine triphosphate (ATP), their relative
effective energy values are not the same as when they are directly incorporated into body fat and body protein.
For an accurate prediction of the feeding value of combinations of feed ingredients, or combinations of
digestible nutrient intakes, computerised mechanistic models that adequately represent nutrient utilisation
should be used. In these models, energy utilisation should be represented based on the partitioning of dietary
nutrients to the various body functions.
4. Significance of microbial fermentation in the upper
gut of pigs
The use of ileal amino acid digestibility is now well established as the method of choice for evaluating the
protein component of pig feeds. The extensive changes to the amino acid composition of the digesta passing
through the large intestine mean that faecal protein is largely microbial with an amino acid composition that is
little influenced by the diet. It is recognised, however, that the pig's microflora are not confined to the large
intestine but that large populations inhabit the upper gastrointestinal tract. Moreover, the ileal microflora is


33
metabolically very active. In consequence, the ileal approach does not completely distinguish between host and
microbial digestion. This is evident from the extent to which NSPs, which are resistant to mammalian digestive
enzymes, are digested before they reach the caecum (Table 12).
Figure 6.2. Table 12. Distribution of NSP fermentation in the gastrointestinal tract of
the pig (Graham s mtsai., 1986; Longland s mtsai., 1990; Zhu s mtsai., 1993; Millard
s Cheeson,1992

Although most NSP digestion occurs in the large intestine, more than one third of it can occur in the upper part
of the gut. This has implications for both carbohydrates and amino acids. The energy losses in the fermentation
of carbohydrates account for as much as one third of their digestible energy. This is not because short-chain
fatty acids (SCFA) are inefficiently utilised but mainly because energy is lost as heat and as combustible gases
(methane and hydrogen). These losses mean that fermented carbohydrates have a net energy value only about
two thirds that of carbohydrates absorbed as simple sugars. In the case of NSPs, these losses are unavoidable,
since it is only by fermentation that their energy can be utilised at all. Starch, however, which represents the
major energy source in pig diets, is potentially recoverable as glucose, provided that it is completely digested by
host enzymes and not by microbial fermentation. Ileal starch digestibility in most pig diets is very high (cereal
starch, for example, is almost completely digested before the end of the small intestine). However, the fact that a
substantial proportion of NSP is also digested before the end of the small intestine makes it probable that some
proportion of starch digestion in the small intestine, especially of the more resistant starches, is achieved by
fermentation. It is probable that much of the starch fermented in the small intestine is converted to lactate.
The implications of pre-caecal microbial activity for amino acids are twofold. First, it means that ileal amino
acid digestibility values for ingredients with substantial amounts of NSP will tend to be in error. If the
microflora have adequate energy, the major end-product of their utilisation of protein, both dietary and
endogenous, in the small intestine is microbial protein, though some ammonia may also be produced. The amino
acids in ileal digesta will thus represent not only undigested food protein and unrecovered endogenous protein
but also microbial protein which will tend to confer its own amino acid pattern on the ileal digesta.
The second implication of pre-caecal microbial activity is that the microflora are synthesising amino acids in a
part of the gut where active amino acid absorption occurs. This means that amino acids synthesised de novo by
the microflora can, in principle, contribute to amino acid supply, augmenting those supplied in the diet. Whether
they do or not depends on the source of nitrogen they use for amino acid synthesis. If they utilise nitrogen from
food protein that would otherwise be absorbed, they simply form an additional step in the utilisation of that food
protein. If, on the other hand, they utilise nitrogen of resistant food proteins that would otherwise escape host
digestion, or if they utilise urea or ammonia to form indispensable amino acids, they make a net contribution to
the amino acid supply of the pig. These questions have yet to be fully answered.
5. Feed ingredient specific effects on nutrient
metabolism in visceral organs
Visceral organs (gastro-intestinal tract, liver, kidney, spleen, pancreas, heart, lungs, reproductive organs)
contribute approximately 15 to 20% of body weight in the growing-finishing pig, yet they are responsible for
close to 50% of total body oxygen consumption. This is equivalent to approximately one third of total energy
intake. Furthermore, more than 50% of whole body protein turnover occurs in these organs. The possibility of
manipulation of energy expenditure and amino acid metabolism in these organs would represent a means to
enhance the efficiency with which dietary nutrients are converted to useful animal products. This possibility is
supported by the large degree of variability in the size and energy expenditure of visceral organs that has been


34
demonstrated between different groups of pigs. This variation may be attributed to differences between pig
genotypes, disease status, feeding levels and diet composition.
Energy expenditure in visceral organs appears to be closely related to the size of the organs. In other words, total
energy expenditure in visceral organs is determined by the number of cells, rather than by energy expenditure
per cell, in these organs. It is thus likely that diet effects (feeding level, specific feedstuffs or nutrients) on
energy expenditure in visceral organs can be predicted reasonably well from diet effects on the size of the
visceral organs.
6. Differences in nutrient digestibilities between
different classes of pig
There is an increasing body of evidence to suggest that there are some important differences in nutrient
digestibilities between different classes of pig. This has important consequences for feed formulation as we
often ignore the effects of body weight on nutrient digestibilities and we extrapolate nutrient digestibilities
obtained from growing pigs to either starter pigs or sows.
Faecal nutrient digestibilities in growing pigs increase with body weight. Furthermore, these changes are related
to dietary fibre levels. These observations indicate that the contribution from hindgut fermentation to faecal
nutrient digestibilities increases with increasing body weight. The effects of body weight on apparent ileal
nutrient digestibilities appear negligible in growing pigs.
Recently, observations on faecal and ileal nutrient digestibilities have been made in sows. In most of these
studies, substantially higher faecal digestibilities have been observed in sows as compared to growing pigs.
Furthermore, the differences in nutrient digestibility between sows and growing pigs vary among ingredients.
The apparent ileal amino acid digestibilities in a range of feed ingredients are higher in sows than in growing
pigs (Table 13.).
Figure 6.3. Table 13. Apparent ileal digestibilities of amino acids in growing pigs and
lactating sows (de Lange and Fuller, 2000)

Again these differences vary with the type of feed ingredient. For example, the differences are larger for corn
and wheat than for most of the other feed ingredients. There is no apparent explanation for this. It may be
attributed to larger retention times of feed in the small intestine, lower endogenous gut amino acid losses, or to a
larger contribution of microbial fermentation to digestion in the small intestine in sows as compared to growing
pigs.


35
Some of these effects of body weight on nutrient digestibility may be attributed to the small negative correlation
between feed intake and nutrient digestibility.
All of these observations suggest that there are effects of body weight and differences between classes of pig
(starter and grower pigs and sows) in both ileal and faecal nutrient digestibilities. These effects and differences
should be considered in feed formulation. It also supports the need to determine nutrient digestibilities in feed
ingredients in all of the various classes of swine.
Questions:
1. How hydrothermic treatments can influence the digestibility of pig diets?
2. Explain the effect of gut microflora on the digestibility of nutrients in pigs!
3. Which way NSP materials can influence the absorption and utilisation of the nutrients of feeds?

36
evaluation for poultry
The distinctive aspects of poultry production that have a bearing on feed evaluation techniques include the
following:
1. The widespread use of uncontrolled, ad libitum feeding places a special emphasis on prehension and feed
intake in the prediction of animal response. In particular the initiation of feeding at day-old is of critical
importance in poultry meat production. As a consequence of this there is a special emphasis on the physical
form of the feed.
2. The digestive tract is relatively short and simple with a rapid rate of feed passage. The lower tract plays a
major role in the maintenance of water balance. As a consequence and in combination with intensive production
methods, this creates a significant emphasis on excreta dry-matter levels. The practical consequence is wet litter
conditions and possible deterioration of carcass quality.
3. The gut microflora play a major role in modifying the response to feed. The consequence is that many feed
additives are used to influence the development of the microflora or to modify the microflora. Important
interactions between feed components are mediated through changes in the microflora. Recent developments in
Europe following the banning of antibiotic growth promoters have tended to emphasize the importance of this
topic.
4. The combined excretion of faeces and urine means that experimental determination of metabolisable nutrients
is simpler than the determination of digestibility. Surgical intervention at higher levels of the tract can overcome
this but these techniques are not very suitable for routine use. A possible exception is the measurement of
nutrient uptake at the terminal ileum in experimental subjects that are sacrificed.
5. Poultry genotypes change very rapidly. As a result there is probably an ever-increasing need for accuracy in
the control of feed composition. Data may require constant re-evaluation.
6. The low margin poultry industry creates, in many countries, the need for very precise control and prediction
of animal response. This places an emphasis on rapid uptake of research, sophisticated databases, quality control
methods and 'real-time' quality control procedures such as NIR.
1. Feed evaluati
2. Energy
The determination and use of metabolisable energy (ME) to control energy content of poultry feedstuffs is still,
more or less, universal. More specifically the apparent ME corrected to zero N-balance (AMEn) is used. The
only fixed factor considered routinely is bird age. There is probably sufficient evidence to justify the routine
distinction between AMEn values for young chicks and adults although the precise demarcation age is not clear.
3. Experimental methods in feed energy evaluation
Energy losses as gas are usually ignored in poultry studies. As defined here, using total excreta energy, the
apparent ME (AME) is being measured. If endogenous energy loss is deducted from the total excreta energy
then true ME (TME) is obtained.
A wide variety of techniques has been described for the experimental determination of energy balance. Direct
measurements of energy intake and loss are widely used but many studies also use digestive markers. Markers
used may include chromium, titanium dioxide, and also acid insoluble ash, although others, such as 'fibre'
fractions, polyethylene and magnesium ferrite, have been reported. Many variations in technique simply reflect
the cost and practical difficulty of carrying out energy balance experiments on a large scale.
An energy balance experiment requires accurate measurement of feed intake and the corresponding excreta.
Errors involved in measuring intake include: feed spillage, loss of feed in the drinkers and changes in feed

Advances in feed evaluation for
poultry

37
composition during the trial (especially dry matter content). Errors in measuring energy output include: loss of
excreta on cages or during collection; loss of volatile components during the experiment or during excreta
drying; failure to establish a constant dry matter for analysis and calculation; failure to balance the gut contents
of the birds at the beginning and end of the balance (end-effects). End-effects can be controlled by having the
digestive tract 'empty' at the start and end of the balance or by making it equally 'full' and using suitably long
collection periods so that any error is relatively small.
Any differences in N-balance between subjects or between treatments in an energy balance experiment will
produce a bias. Thus the convention has arisen that all balance data are corrected to a zero-N balance basis. A
constant related to the excretion of uric acid is used for the energy value corresponding to excreted nitrogen -
frequently 34.4 kJ/g.
The relationships involved in ME determination are shown in Figure 5. shows the supposed linear relationship
between feed intake and excreta energy output. Endogenous loss is estimated by the intercept on the y-axis of
this graph. TME is proportional to (1-b) where b is the slope of the line. AME is proportional, in the same way
but for a given intake, to the slope of a line joining a point on the excreta output line and the origin. As a
consequence AME clearly varies with feed intake (as shown in Figure 5.) except in the special case where
endogenous loss is zero and TME = AME. The y-axis in this graph may be either total excreted energy or
energy excretion corrected for N-balance as described above.
Figure 7.1. Figure 5. The relationship between feed intake and energy excretion and
different ME values (Fisher, 2000)

In 1976 Dr Ian Sibbald from Canada (Sibbald 1976) proposed an assay for ME measurement based on placing a
small and known aliquot of feed directly into the crop of a starved ('empty') cockerel. This avoided the need to
measure intake and avoided practical sources of loss such as feed spillage. By using a starved subject a short
assay period could be used, all the energy containing materials being excreted after 24 to 48 hours post-feeding.
The method required only small amounts of feed and feed ingredients could be used without incorporation in a
complete feed. Thus the statistical errors of feed substitution methods were also avoided. This procedure had
advantages from a logistical point of view but the small dose used also meant that, relative to total intake,
endogenous loss of energy was a significant proportion of the total balance. Thus determination of the
endogenous energy loss was necessary, together with correction of the measured AME to give a TME value.
The method became widely, but erroneously, known as the TME method. The small dose also means that any
errors are large relative to the total energy balance and a scrupulous technique is required.
From an experimental point of view three basic types of ME assay still have a valid role in different
circumstances. It is scarcely relevant to ask which of these is the 'best'!
1. Intubation of small amounts of unmixed ingredients into the crop of starved subjects, probably adult roosters,
combined with total excreta collection. This will be close to the original Sibbald (1976) proposal with some
modification of timing, use of glucose feeding to reduce energy loss and practical issues, which improve
accuracy (see McNab (1990) for a discussion). This method should be chosen when the logistical and cost
advantages of small-scale work are paramount.
2. So-called 'rapid' assays, which combine the use of a starved or 'empty' bird with free (or nearly free) feeding
to give normal intakes (see Farrell 1981) for a discussion). These methods were designed to avoid the
controversies about force-feeding and about using small amounts of single feeds. The logistical and cost
advantages may be retained but, in most cases, ingredient substitution methods must be used.

poultry

38
3. The use of a larger traditional assay using'full' subjects with free feeding. Special attention is paid to
eliminating end-effects and to avoiding variation in food intake. From a logistical point of view these
experiments are much more complex and costly. Ingredient substitution methods must normally be used.
4. Prediction of the ME values of poultry feeds and
feedstuffs
Prediction of energy values may be based on chemical analysis or in vitro measurements or on a combination of
these. Equations may be completely empirical or include mechanistic concepts. Many examples for individual
classes of feedstuffs can be found in the literature. Here some general points may be noted. Firstly, it has to be
clear that the prediction of energy values of individual feedstuffs and for mixed feeds present completely
different problems. For mixed feeds the digestibility of nutrients will vary within a fairly small range and
chemical predictors are surprisingly successful. The digestibility of nutrients in feedstuffs will vary
systematically and, therefore, prediction methods that can be applied across classes of feedstuffs will have to
consider this, as well as chemical composition, in some way. It may well be that successful equations for
ingredients can only be developed within classes rather than for general application.
For complete feeds, but not individual ingredients, simple chemical prediction equations have proved to be
surprisingly robust. The most widely used, the so-called 'EEC' equation, uses fat, protein, starch and sugars as
additive predictors.
The EEC equation is as follows:
AMEn (MJ/kg) = 0.3431%FAT + 0.1551%CPR + 0.1669%STC + 0.1301%SUG
where:
FAT = crude fat determined after acid hydrolysis
CPR = crude protein
STC = starch determined by polarimetry
SUG = free sugars determined by Luff-Schoorl method.
5. Prediction equations for feedstuffs
The European Table of Energy Values For Poultry Feedstuffs (1989) has been published by the European
Federation of the Worlds Poultry Science Association. The table contains information mean AMEn values for
each product, the chemical composition of a sample which corresponds to this mean value, calculation factors
for adjusting the AMEn value of samples with different composition and averages digestibility coefficients for
crude protein, crude fat and NFE. According to this table three different methods are suggested for predicting
the AMEn values of feedstuffs. These are the regression lines, equation lines and using digestibility coefficients.
Multiple regression equations are suggested for feedstuffs when the ME could be predicted significantly from
the nutrient composition of several samples. The calculation factors are the dry matter, ash, crude protein, crude
fat, crude fibre and starch content.
Equation lines are used for some feedstuffs with extreme composition (groundnut, cottonseed sunflower
products). The calculation factors are dry matter, crude fibre and fat. For the calculation of AMEn from
digestible nutrients the following coefficients were used: digestible fat: 38.8 KJ/g; digestible crude protein:
18.03 KJ/g; digestible NFE: 17.32 KJ/g.
6. Prediction equations for feed fats
The energy values of fats are notoriously variable and difficult to deal with in practice. Fat 'quality' is also a
wider topic, involving impurities and toxins as well as energy values. Apart from linoleic acid, specific fatty
acid compositions are not usually controlled in poultry feed formulation but knowledge of the total composition
of fat is still required.

poultry

39
From an experimental point of view, fats present particular problems. Firstly, they can only be included in the
feed at quite low levels. Thus large errors of extrapolation, inherent in substitution methods, are inescapable.
Secondly, fats show synergistic effects on digestibility, thus the ME values of added fat and of the basal feed are
not additive. Thirdly, the level of fat inclusion itself affects digestibility and hence measured ME. Finally, there
are interactions between dietary fats and other ingredients and nutrients.
In spite of much research, a complete answer to the problems of evaluating fats is not available today. The most
fundamental remaining problem is that fat feeding values (energy) are not additive because of synergistic effects
and thus the basic conditions of linear programming are not fulfilled.
7. Interactions among energy yielding ingredients
The basic assumption of both energy evaluation experiments and commercial feed formulation is that the values
ascribed to individual ingredients are additive when used to calculate the energy content of a mixture.
Some cases of non-additivity are well known. As well known examples the synergistic effects amongst fats of
different compositions and the effects that tannins have on digestibility of all components of the feed may be
quoted.
Research on variation in the ME values of wheat led, eventually, to recognition of the general role that dietary
non-starch polysaccharides (NSPs) may play by interfering with nutrient absorption. The most potent
interaction, and probably the one of most practical importance, occurs between NSPs and lipid digestibility.
This can be seen clearly in the effects of basal diets containing wheat on the determined ME values of fats.
Ingestion of NSPs increases the viscosity of gut contents and increases moisture and bacterial colonisation. The
raised viscosity of the gut content increases retention time of the digesta, increases the thickness of the unstirred
water layer adjacent to the mucosa and reduces diffusive and convective transport. Collectively these changes
result in reduced solubilisation of fat and (possibly) hydrolysis of fat and protein. Nutrient uptake at the mucosal
surface is reduced and uptake is also reduced by the effects of enhanced bacterial growth on the proliferation
rate of enterocytes and changes in the morphology of the vilii and microvilli. Microbial activity increases the
deconjugation and loss of bile acids, impairing the return of bile acids to the liver and subsequent recycling.
Finally endogenous nutrient losses may be enhanced by digestion of viscous NSPs. The incorporation of these
important issues into routine feed formulation has not been completely solved at this time.
8. Enhancing dietary energy values
Enhancement of feed energy values is possible by the addition of feed additives and by feed processing.
9. Feed enzymes
Feed enzymes, especially carbohydrases, now play an essential role in the feeding of many classes of livestock.
Their use in wheat and barley based feeds for broiler and turkeys is more-or-less universal. Commercial
development of enzymes for maize-soya bean based feeds is now the centre of activity.
Enzymes have a number of claimed values but, in the context of feed evaluation, it is the enhanced digestibility
of energy and amino acids that predominates. Fat digestibility is also noticeably enhanced in wheat-based feeds
reflecting the effect of the enzymes on the viscosity of gut contents. Enzymes also probably encourage the
development of a more benign gut microflora.
The undoubted effects of feed enzymes can be taken into account in practical feed evaluation in a number of
ways. All formulation data can be left unchanged except for reducing the formulated level of energy.
Alternatively the energy (and other) values can be increased for some ingredients. The final alternative is to
ascribe all of the effects to the enzyme premix and to give this a nutritional value that reflects the changes in all
ingredients.
Of these, the second possibility seems to be the most sound because it allows differences in the effect of
enzymes on different ingredients to be taken into account. The use of enzymes in a programme of feed
evaluation experimentation obviously needs to be carefully considered.

poultry

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10. Protein and amino acids
Controlling amino acid and nitrogen supply to poultry is more straightforward than for energy. There are few
completely new developments but more a gradual improvement in concepts and in the application of existing
data. A good commercial scheme today would probably contain the following elements:
- Classification of ingredients by process, source etc.
- Total amino acid compositions covering 5-7 amino acids.
- Adjustment of total amino acids for protein content using prediction equations or in proportion to protein
content.
- Control, in feed formulation, of digestible amino acid and crude protein levels.
The greatest weaknesses in such a scheme arise from four main sources. First of all there are continuing
difficulties over amino acid analysis. Secondly, availability coefficients are presumably variable within an
ingredient but we have no methods of predicting such variability. Thirdly, there is a well-recognised need to
move from the measurement of availability in excreta to techniques for assessing availability at the ileum.
Fourthly, control of total protein level or non-essential amino acids is not fully considered.
11. Measurement of amino acid availability
As noted above, most existing data available for practical application at this time are based on excreta analysis.
Corrections may or may not be made for endogenous amino acid loss to give values for true or apparent faecal
digestibility. It is clearly important for users to know which they are using when the general term 'digestibility'
is used.
12. Feed formulation and the control of protein and
amino acid levels
There is a range of additional topics in protein nutrition of poultry, which collectively, limit our ability to
describe feedstuffs as sources of protein and to accurately predict bird performance. These may be described as
the limits to accuracy in protein nutrition.
Up to four essential amino acids are available in crystalline form at feed-grade prices, methionine, lysine,
threonine and tryptophan. When these are used, and to the extent that they are limiting, then protein mixtures,
which are perfectly balanced with respect to these amino acids, may be devised. The controls placed on other
amino acids, which are supplied only from intact proteins, will then determine the total crude protein content of
the feed. The simplest case is that the minimum protein level will be determined by the level of the amino acid
which is next limiting after those supplied in pure form. Protein is often controlled at higher levels than this, for
commercial reasons.
Crystalline amino acids are used increasingly to meet the requirements of poultry. Initially this is on economic
grounds but increasingly their use is being encouraged by concerns over N-pollution.
13. Information sources for amino acids and protein
digestibility
The European Table of Energy Values for Poultry Feedstuffs (WPSA 1989) contains figures for protein
digestibility.
Results of amino acid analysis published by Degussa AG are perhaps the most complete data set on this topic.
These tables also include prediction equations relating amino acid levels to crude protein content and
availability coefficients based on true faecal digestibility. The tables are available as a database for computer use
and calculation. The other amino acid companies (Eurolysine, ADM, Rhone Poulenc Nutrition etc.) also publish
good tables including data on availability. In 1992 the WPSA published The European Amino Acid Table. This

poultry

41
contains average results of amino acid analyses collected at various centers in Europe. However it is of limited
value as the important question of variation in amino acid analysis is not addressed.
14. Minerals
Whilst most emphasis in feed evaluation for poultry is placed on energy and protein, evaluation of some mineral
sources has been important in recent years. Recent work on phosphorus availability has been driven, especially
in the Netherlands, by concerns about pollution. This creates a need for very precise methods of control in
animal feeding.
For the same reason, phytase enzymes are now in widespread use. These increase the utilisation of phytate-P
(inositol-bound P), permitting an overall reduction in total P levels in the feed. The following issues arise in this
field:
- Poultry do not secrete phytase digestive enzymes. Thus any degradation of phytase depends on bacterial
phytase in the gut or on endogenous phytase in the feed.
- Endogenous phytase is found in some ingredients, especially wheat. Phytase, either endogenous or exogenous
is very heat labile, and therefore, in principle, different availability figures should be used for ingredients used in
mash or pelleted feeds.
- In recent years, at least in Europe, the move to using all-vegetable feeds more widely has meant that phytate
levels in feeds have tended to increase.
- In vegetable feeds the calculation of availability in mineral phosphates becomes very important. This is a
difficult area but it appears that some progress can be made by calculating digestibility from the mix of chemical
forms that is found in different products. Even products with the same name, for example dicalcium phosphate,
may differ widely in the actual chemical forms present. The chemistry of these products is not precisely known
however.
Four approaches to the evaluation of P-availability are used:
1. The classical approach based on the simple hypothesis that P from all plant sources is 1/3 available whilst P
from animal and mineral sources is completely available.
2. Systems based on the experimental determination of P-availability using chick toe-ash as the assay response.
3. Systems based on the experimental determination of P-availability using phosphorus balance (metabolisable P
or retained P).
4. Chemical based systems using the analysis of phytate-P.
Questions:
1. What is the difference between AME and TME? Describe the techniques of ME determination!
2. What are the advantages to compose poultry diets based on digestible amino acid levels?
3. What kind of methods can be used for the determination of endogenous amino acid losses?
4. What are the main advantages of using available phosphorous in the diets formulation?

42
evaluation for ruminants
1. Energy evaluation systems
1.1. Early energy systems
Energy systems have a history going back to the first half of the nineteenth century but did not provide an
adequate description of energy utilisation until methods of animal calorimetry came into use in the second half
of that century. About 1900, H. P. Armsby at the University of Pennsylvania and O. Kellner at the Mckern
Experiment Station in Germany used the results of their calorimetric studies to devise energy systems based on
the net energy value of foods. The systems differed in some respects, most evidently in the units used. Armsby
expressed net energy in terms of calories (the unit antedating joules), but Kellner - believing that farmers
would have difficulty in understanding calories - expressed net energy values of foods relative to the net energy
value of that common food constituent, starch. For example, if the net energy value of barley was found to be
1.91 Mcal (megacalories) per kilogram, and that of starch, 2.36 Mcal/kg, then 1 kg of barley was stated to have
a starch equivalent of 1.91/2.36 = 0.81 kg. Both Kellner's and Armsby's systems encountered difficulties caused
by the differences in net energy values of foods for maintenance, growth, etc. and used approximations to
avoid these difficulties. Kellner's starch equivalent system was used (mainly in Europe) as the basis of practical
rationing systems until the 1970s.
Armsby's net energy system was incorporated in what was at one time the standard reference work on the
feeding of livestock in the USA, F B Morrison's Feeds and Feeding, but was not much used in practice. The
preferred system in the Americas was for many years the total digestible nutrients (TDN) system.
1.2. Recent systems
All three components of a modern energy system, the energy content of foods, the energy requirements of
animals and the interface linking them, can be given greater accuracy by introducing additional factors.
As the food intake of an animal increases, the metabolisability of the food energy declines. Food intake can be
defined as the level of feeding, which is the metabolisable energy intake relative to that required for
maintenance; thus if the level of feeding is 60/33=1.82. For growing cattle the level of feeding is commonly 2-
2.5, but for lactating cattle it rises to 3-4. The energy requirement of lactating cows is increased by 1.8 per cent
for each unit increase in the level of feeding. Thus for a cow with a level of feeding of 3 the initial estimate of
energy requirement would be increased by 2 x 1.8 = 3.6 per cent to allow for the effects of increasing food
intake on the energy content of the ration. The same correction is made for lactating ewes, but not for other
classes of ruminants.
Increasing the level of feeding may also reduce the efficiency of utilisation of ME (i.e. reduce the k factors).
Refinements of the system include corrections for this effect. For example, for growing cattle kg is assumed to
have its normal predicted value when the level of feeding is at twice the maintenance level, but if the level of
feeding is greater than this, kg is reduced, and if the level of feeding is less than twice the maintenance level, kg
is increased. For example, if cattle on a diet containing 10 MJ ME/kg DM were fed at 2.5 times maintenance, kg
would be reduced from 0.43 to 0.39. The same correction is used for growing lambs.
There is also evidence that when metabolisable energy is used for growth, the efficiency with which it is utilized
varies with the nature of the diet. For example, when diets containing 11 MJ ME/kg DM are made up from
either high-quality forage alone or poorer-quality forage plus concentrates, the diet containing concentrates will
have a kg value about 5 per cent higher than that of the diet of forage alone. Among the forages, there is
evidence that kg values are greater for first growths (i.e. spring growths) of temperate herbages than for later
growths having the same metabolisable energy concentration, and are higher for temperate than for tropical
forages. However, it is difficult to classify diets into different categories for the purpose of predicting kg.
Although it seems undesirable to introduce an arbitrary correction factor to an otherwise logical system, it is
important that the system should accurately predict the animal growth rates that are achieved in practice.

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43
In 1988 the European Association of Animal Production carried out a survey of the feeding systems in use in
European countries. The report on energy systems for ruminants demonstrated the wide variety of systems in
use in Europe, and it is not possible to describe all of them here. The Netherlands, Belgium, France, Germany,
Switzerland, Italy and Austria have systems with many features in common, and the Dutch system will be
described as an example. The metabolisable energy content of foods is calculated from digestible nutrients and
is then converted to a net energy value. For growing animals, the basis for this conversion is that the animal
production level is assumed to be constant, at 1.5, and hence that kmp, has a unique value for a food of known
metabolisable energy concentration. Each food can therefore be given a single net energy value for maintenance
and production (NEmp), but this is converted to a unit value by dividing it by the presumed NEmp of barley (6.9
MJ/kg, or about 8 MJ/kg DM).
For lactating cows a corresponding net energy value for maintenance and lactation is calculated by assuming
that kl is 0.62 and the net energy value of the food for lactation (called NEl) can also be calculated.
After using a system of total digestible nutrients for many years, the USA changed to net energy systems for
beef and dairy cattle, these being described in publications of the National Research Council. The Hungarian
system is almost the same. Metabolisable energy is calculated as 0.82 x digestible energy and digestible energy
is calculated as 18.45 MJ per kg TDN. For beef cattle, foods are given two net energy values, for maintenance
(NEm) and gain (NEg), these being calculated from the metabolisable energy (ME) content of dry matter in
each food by means of the following equations:
NEm = 1.37ME - 0.033ME2 + 0.0006ME3 4.684
NEg = 1.42ME - 0.0416ME2 + 0.0007ME3 6.904
Where: ME = metabolisable energy content of dry matter, and all energy values are expressed in MJ/kg.
Net energy values for lactation (NEl) for the US system have been calculated from TDN, digestible energy by
the following equation, where df (discount factor) is correction factor related to the fibre content of fedstuffs :
In the future, as energy systems are modified to incorporate new findings they are likely to become even more
complex. As the need for simplicity of calculation is diminished by the increasing availability of computers,
energy systems are becoming parts of much larger mathematical models of nutrient requirements that are
capable of dealing simultaneously with energy, amino acids, vitamins and minerals (and with the interactions
between them). Complex systems tend to obscure the principles upon which they are based. Students of animal
nutrition should therefore pay particular attention to the principles of energy metabolism outlined in the previous
chapter and at the same time should familiarise themselves with the energy systems currently used in their own
countries.
The use of DCP for evaluating food proteins for ruminants has been largely abandoned. This resulted from a
growing awareness of the extensive degradative and synthetic activities of the microorganisms of the rumen.
Rumen microorganisms are responsible for providing the major part of the energy requirements of the host
animal by transforming dietary carbohydrates to acetate, propionate and butyrate. In order to do this and to
exploit the energy potential of the food fully, they must grow and multiply and this involves large-scale
synthesis of microbial protein. The nitrogen for this is obtained, in the form of amino acids, peptides and
ammonia, by breakdown of the nitrogen fraction of the food. Bacteria acting on the structural carbohydrate (SC)
fraction of the diet use only ammonia, whereas those acting on the non-structural fraction (NSC) derive about 65
per cent of their nitrogen from amino acids and peptides, and the remainder from ammonia.
The microbial protein passes from the rumen, is digested in the small intestine, and so makes a contribution to
satisfying the nitrogen requirements of the host animal. The magnitude of this contribution depends upon the
speed and extent of microbial breakdown of the dietary nitrogen fraction, upon the efficiency of the
transformation of the degraded material into microbial protein (nitrogenous compounds), the digestibility of the
microbial protein and the biological value of the latter.
The degradative and synthetic processes taking place in the rumen are of major importance in the nitrogen
economy of the host animal since they determine the nature of the amino acid mix made available for protein
synthesis at tissue level. Satisfying the demands of the rumen microorganisms for readily available nitrogen is
a major function of the diet and to this end a certain proportion of the nitrogen fraction must be degradable by
the rumen microorganisms.

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Current systems for the evaluation of food protein for ruminant animals involve determinations of the
degradability of protein in the rumen, the synthesis of microbial protein, the digestion in the lower gut of both
food and microbial proteins, and the efficiency of utilisation of absorbed amino acids. The methods used to
determine these components of the system are described next, after which their use in the systems will be
illustrated.
2. Protein evaluation for ruminant animals
Traditionally, proteins in foods for ruminant animals have been evaluated in terms of crude protein (CP) or
digestible crude protein (DCP). Realization that the crude protein fraction contained variable amounts of non-
protein nitrogen led to the use of true protein instead of crude protein but this was unsatisfactory since no
allowance was made for the nutritive value of the non-protein nitrogen fraction. The concept of protein
equivalent (PE), introduced in 1925 but now no longer used in this context, was an attempt to overcome this
difficulty by allowing the non-protein nitrogen fraction half the nutritive value of the true protein. The term
`protein equivalent' is currently used in connection with foods containing urea. Such foods must by law be sold
with a statement of their content of protein equivalent of urea. This means the amount of urea nitrogen
multiplied by 6.25.
The use of DCP for evaluating food proteins for ruminants has been largely abandoned. This resulted from a
growing awareness of the extensive degradative and synthetic activities of the microorganisms of the rumen.
Rumen microorganisms are responsible for providing the major part of the energy requirements of the host
animal by transforming dietary carbohydrates to acetate, propionate and butyrate. In order to do this and to
exploit the energy potential of the food fully, they must grow and multiply and this involves large-scale
synthesis of microbial protein. The nitrogen for this is obtained, in the form of amino acids, peptides and
ammonia, by breakdown of the nitrogen fraction of the food. Bacteria acting on the structural carbohydrate (SC)
fraction of the diet use only ammonia, whereas those acting on the non-structural fraction (NSC) derive about 65
per cent of their nitrogen from amino acids and peptides, and the remainder from ammonia.
The microbial protein passes from the rumen, is digested in the small intestine, and so makes a contribution to
satisfying the nitrogen requirements of the host animal. The magnitude of this contribution depends upon the
speed and extent of microbial breakdown of the dietary nitrogen fraction, upon the efficiency of the
transformation of the degraded material into microbial protein (nitrogenous compounds), the digestibility of the
microbial protein and the biological value of the latter.
The degradative and synthetic processes taking place in the rumen are of major importance in the nitrogen
economy of the host animal since they determine the nature of the amino acid mix made available for protein
synthesis at tissue level. Satisfying the demands of the rumen microorganisms for readily available nitrogen is
a major function of the diet and to this end a certain proportion of the nitrogen fraction must be degradable by
the rumen microorganisms.
Current systems for the evaluation of food protein for ruminant animals involve determinations of the
degradability of protein in the rumen, the synthesis of microbial protein, the digestion in the lower gut of both
food and microbial proteins, and the efficiency of utilisation of absorbed amino acids. The methods used to
determine these components of the system are described next, after which their use in the systems will be
illustrated.
3. Degradability of the nitrogen fraction of the diet
Nitrogen fractions within the diet will vary in their susceptibility to down, from immediately degraded to
undegradable, and from 0 to 1 extent to which they are degraded in the rumen and digested when they reach the
small intestine.
Degradability will be affected by such factors as the surface area available for microbial attack and the
protective action of other constituents a as the physical and chemical nature of the protein. Claims have been
that the solubility of a protein is correlated with ease of breakdown but these do not survive critical examination.
Thus casein, which is degraded in the rumen, is not readily soluble; whereas albumin, which is resistant to
breakdown, is readily soluble. It has been suggested major factor affecting degradability is the amino acid
sequence within the protein molecule. If this is so then the nature of the microbially rumen peptidases is of
considerable importance and it seems whether any simple laboratory test for degradability is possible.

ruminants

45
The extent to which a nitrogen fraction is degraded in the rumen will depend upon its innate degradability and
the time it spends in the rumen, i.e. upon rate of passage. As the rate of passage increases so the extent of
ruminal breakdown is reduced.
3.1. Measurement of degradability in vivo
This involves the measurement of dietary nitrogen intake, the endogenous nitrogen (EN), the non-ammonia
nitrogen (NAN) and microbial nitrogen (MN) of dietary origin passing the duodenum. Degradability (Dy) of
nitrogen is then expressed as:

This method requires accurate measurement of duodenal flow and microbial and endogenous nitrogen. The
flow measurement, which requires the use of a dual-phase marker system, has a large coefficient of variation
(between animals) and many published values must be suspect owing to the small number of animals used in
their determination. Microbial nitrogen in duodenal nitrogen is usually identified by means of marker substances
such as diaminopimelic acid (DAPA), aminoethylphosphoric acid (AEPA), ribonucleic acid and amino acids
labelled with 35S, 32P and 15N. The concentration of marker in the microorganisms is measured in a sample of
rumen fluid. Different markers may give results which vary widely, sometimes by as much as 100 per cent. The
assumption that the microorganisms isolated from rumen fluid are representative of those in the duodenum is of
doubtful validity, since the latter include organisms normally adherent to food particles and/or the rumen
epithelium. The endogenous fraction constitutes about 50-200 g/kg of the duodenal nitrogen but is difficult to
quantify. A value of 150 g/kg is frequently assumed. Measurements of degradability are thus subject to possible
errors owing to uncertainties in measuring duodenal flow and microbial endogenous nitrogen, and are affected
by dietary considerations such as of feeding and the size and frequency of meals. It has been calculated that
estimates of degradability may vary over a range of 0.3-0.35 owing to errors of determination alone. Despite its
inadequacies this technique remains the method currently available for providing an absolute measure of protein
degradability and is the standard against which other methods have to be assessed.
3.2. Determination of degradability in sacco (or in situ)
This involves incubation of the food in synthetic fibre bags suspended in the rumen. The degradability is
calculated as difference between the nitrogen initially present in the bag and that present after incubation, stated
as a proportion of the initial nitrogen. When protein disappearance (p) is regressed on time, p increases a
reducing rate. The relationship may be described by the curve, illustrated in Fig. 6..
Figure 8.1. Fig 6. Relationship of protein disappearance to time of incubation

The technique is subject to several inherent sources of error which must be controlled if reproducible results are
to be obtained. Chief of these are sample size, bag size, porosity of the bag material and treatment of the bags
following removal from the rumen. Ring tests (i.e. tests carried out at several laboratories) have shown
unacceptably high inter-laboratory variability, indicating a need for strictly defined standard procedures which
have to be rigidly adhered to if the results are to be universally applied in practice.
A basic assumption of this method is that disappearance of nitrogen from the bag, virtually reflecting solubility
in rumen fluid, is synonymous with degradability. It has been known for some time that small amounts of food
protein which are solubilised may leave the rumen without being degraded, and this must cast doubt on the
veracity of values obtained using the technique. Even more serious in this context is the recent observation that
acid-detergent insoluble nitrogen (ADIN), known to be undegradable, may disappear during incubation. A
further complication is the presence in the bags of rumen bacteria, which contribute to the nitrogen of the
contents.

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46
4. Laboratory procedures for determining nitrogen
degradability
4.1. Solubility in buffer solutions
Significant correlations have been demonstrated between the de values for the nitrogen fractions of foods and
their solubility characteristics in a range of buffer solutions, including McDougall artificial saliva, borate
phosphate buffer and Wise Burroughs's buffer. When the methods are used for a range of foods, errors of
prediction may be high but within food types predictions are improved sufficiently to allow the use of buffer
solubility in the routine monitoring of concentrate foods.
When used in conjunction with methods for the fractionation of nitrogen to predict degradability, solubility in
buffer solution has good correlation with figures based on enzyme solubility.
4.2. Solubility in enzyme solutions
Solubilisation of protein by purified enzymes from fungi and bacteria has been widely investigated as a means
of estimating degradability. Different proteases have given varying results when compared with the in sacco
technique. This is not unexpected in view of the fact that a single enzyme is being used to simulate the action of
the complex multi-enzyme system of the rumen. As with the buffer solutions, accuracy of predictor is over a
range of foods but improves when the technique is applied within food types.
4.3. Near-infrared reflectance spectroscopy (NIRS)
NIRS measurements reflect the types and proportions of organic structures within a material. As such they are
widely used for the routine analysis of foods and their nutritional evaluation. The technique might therefore be
expected to provide a solution to the problem of determining degradability. Current indications are that NIRS
has the ability to estimate nitrogen degradability with a high degree of precision and considerable accuracy with
r2 values of 0.80-0.87 being claimed.
4.4. Rate of passage
The extent to which a protein is broken down in the rumen depends not only on its innate degradability but also
upon the length of time for which it is exposed to breakdown and therefore upon its rate of passage through the
rumen. The rate of passage of food from the rumen is affected by a complex of food and animal factors. Passage
is faster for:
smaller particles,
particles of higher density,
more highly hydrated particles,
more highly digested particles.
It will thus increase as digestion and rumination proceeds. Rate of passage increases with increased dry matter
intake and is by affected by a number of animal and environmental factors:
advancing pregnancy limits rumen fill and increases rate of passage,
lactation increases intake and rate of passage,
excessive body condition can reduce intake and rate of passage,
high environmental temperatures will reduce intake and throughput.
The rate of passage may be determined by treatment of the protein with dichromate. The treatment renders the
protein completely indigestible, there is no loss of chromium from the protein subsequent to treatment and
particle distribution is not affected. The rate of dilution of chromium in samples of rumen contents taken over a

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47
period of time can therefore vide an estimate of the rate of passage of the protein from the rumen. The rate of
passage of forages may be estimated from the dry matter intake (DMI) of the animal and the proportion of this
provided by forage.
4.5. Efficiency of nitrogen capture
The efficiency with which nitrogen is captured by the microorganisms of the rumen depends not only upon the
speed and extent of breakdown but also upon the synchronous provision of a readily available, utilisable source
of energy to fuel the synthesis of microbial protein. Failure to achieve this balance can result in too rapid and
extensive a breakdown, and the synthetic powers of the rumen microorganisms may be overwhelmed. Wastage
may then occur since the excess ammonia is absorbed and largely excreted as urea; some, though, is recycled
via the rumen wall and contributes further to the nitrogen economy of the rumen. The extent of recycling has
been estimated as about 70 per cent of nitrogen intake for diets of low protein content (about 5 g/kg) and as little
as 11 per cent for foods with about 20 g/kg.
That part of the food crude protein which is immediately degradable is unlikely to be as effective a source of
nitrogen for the microorganisms as that which is more slowly degraded. It is generally considered that the
slowly degraded nitrogen fraction is incorporated into microbial protein with an efficiency of 1.0, whereas that
immediately degraded is less efficiently used. Estimates of the efficiency with which immediately degraded
protein is incorporated vary, but 0.8 is a commonly used figure.
4.6. Yield of microbial protein
The yield of microbial protein which becomes available for digestion and absorption post-ruminally by the host
has been related to the energy of the diet stated in terms of digestible organic matter (DOM), digestible organic
matter digested in the rumen (DOMADR), total digestible nutrients (TDN), net energy for lactation (NEL),
metabolisable energy (ME), fermentable organic matter (FOM), fermentable metabolisable energy (FME) and
rumen-degradable carbohydrate. The last three eliminate fat and products of fermentation, neither of which are
considered to provide energy which can be utilised by the rumen microorganisms. The energy of fermentation
products is significant in the case of silages and some distillery and brewery by-products. Hays are not
considered to have und gone fermentation though they often contain measurable amount fermentation acids such
as acetic and propionic. The routine measure of the contribution of fermentation products to metabolisable
energy individual foods has not been a feasible proposition and assumed values are commonly used.
Sophisticated models attempt to relate microbial yield to the rate carbohydrate fermentation and rate of passage,
theoretical growth the energy cost of bacterial maintenance and the form of nitrogen available to the rumen
microorganisms. Many of the relationships involved such calculations are based on laboratory characterisation
of the food, and the value of the model will depend on the validity of the relationships between the laboratory
determinations and the values used in models.
4.7. True digestibility of protein
The microbial protein synthesised in the rumen may be protozoal or bacterial, the relative proportions
depending upon conditions within that organ. Thus low rumen pH tends to reduce protozoal activity and
stimulate that of certain bacteria. The mixture of bacterial and protozoal protein, along with dietary protein
not degraded in the rumen, passes to the abomasum and small intestine. Here it is broken down to amino acids,
which are then absorbed into the body. The digestibility of bacterial protein is lower (about 0.75) than that of
the protozoal (about 0.90) and the overall digestibility of microbial protein will depend to some extent upon the
rumen environment. However, protozoal protein constitutes some 5 to 15 per cent only of the total microbial
protein flow from the rumen and its influence on the overall digestibility of microbial protein will be small. The
composition of bacteria is variable but that shown in Table 14. is an acceptable approximation.
Figure 8.2. Table 14. Composition of rumen bacteria (g/kg dry matter)

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48
About 15 per cent of the total nitrogen is in the form of nucleic acids, about 25 per cent is cell wall protein and
the remainder is true protein. Available evidence indicates that the digestibility of nucleic acid nitrogen is of the
order of 0.8-0.9, that of microbial true protein 0.85-0.9. It is commonly assumed that the protein associated with
the cell walls is completely indigestible. Most estimates of the true digestibility of microbial protein are of the
order of 0.85-0.87, which is higher than might be expected in view of the proportions of the fractions present in
the protein. Although nucleic acids are highly digestible their nitrogen is of no use to the animal, as after
absorption they are totally excreted in the urine.
The digestibility of the undegraded dietary protein is a characteristic of the protein mix in the food and may vary
considerably from diet to diet. The true digestibility of the undegraded dietary protein will vary with the
proportion of the various protein fractions present. Thus amino acids, peptides, globulins, albumins and
glutelins will be almost completely digested; prolamins, proteins associated with the cell walls and denatured
proteins will have digestibility values of about 0.8; the protein of Maillard products and nitrogen bound to lignin
will be completely indigestible.
Digestibility has been shown to be inversely related to the content of acid detergent insoluble nitrogen (ADIN),
which reflects that part of the food nitrogen which is closely bound to insoluble fibre. The digestible
undegradable protein content (DUP) of a food is calculated thus:
DUP = 0.9 (undegradable protein - ADIN x 6.25)
This equation is based on the assumptions that ADIN is indigestible and that the digestible fraction has a true
digestibility of 0.9.
In the case of foods such as maize gluten and some distillery and brewery by-products, which have been heat
treated under moist conditions, Maillard-type reactions may occur, resulting in an increase the concentration of
nitrogenous compounds insoluble in acid-detergent. Such 'acquired ADIN' does have a finite though low
digestibility and the above equation is unreliable when used for such foods.
4.8. Efficiency of utilisation of absorbed amino acids
The mixture of amino acids of dietary origin absorbed from the small tine (i.e. the truly digested amino acids) is
utilised for the synthesis of tissue protein. The efficiency of this process, which depends upon composition of
the mix relative to that of the protein to be synthesised, is best represented by its true biological value. This
will in turn depend upon the biological values of the digested undegraded dietary protein and digested microbial
protein, and upon the relative proportions of each contributing to the mix. In addition, it will vary with the
primary function which it is required. Microbial protein is thought to have a relative constant biological value of
about 0.8, whereas that of dietary origin variable and characteristic of the foods making up the diet. Prediction
of such dietary values is extremely difficult, since the biological values of the individual proteins are no guide to
their value in combinations.
An alternative approach is to estimate the supply of essential amino acids made available to the tissues (i.e.
those absorbed from the small intestine) and to relate this to the amino acid requirements of the animal. This
approach needs information on the truly digestible amino acid content of the undegraded dietary and the
microbial protein.
The essential amino acid content of ruminal microbial protein is frequently claimed to be relatively constant.
In fact, large differences in the amino acid composition of samples of microbial protein have been shown to
exist (Table 15.).
There is evidence that the essential amino acid composition of undegradable dietary protein may differ
significantly from that of the original dietary material and it has been suggested that estimates of its contribution
to the amino acid supply should be based on the amino acid profile of the insoluble fraction of the dietary
protein rather than that of the whole dietary protein. A comparison of the essential amino acid profiles of whole
and insoluble proteins in some common foods is shown in Table 16.
Figure 8.3. Table 15. Amino acid composition of ruminal bacteria (g/100g of amino
acids)

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49

Figure 8.4. Table 16. Amino acid composition of whole and insoluble protein in some
common foods (g/100g protein)

4.9. The Hungarian metabolisable protein system
The new protein evaluation system was introduced in 1999. Metabolisable protein content of the feed means the
ratio of amino acids absorbed in the small intestine of the animals. These amino acids can originate from the
undegradable protein (UDP) and partly from the rumen degradable protein (RDP) contents of the feeds.
All the feedstuffs have two metabolisable protein contents, according to the potential of the certain feedstuff
supporting the microbial protein fermentation in the rumen:
Energy dependent metabolisable protein (MFE): digestible undegradable protein + potential digestible microbial
protein according to the energy content of the feedstuff
Nitrogen dependent metabolisable protein (MFN): digestible undegradable protein + potential digestible
microbial protein according to the nitrogen content of the feedstuff
digestible UDP = 0.9 (UDP ADIN x 6.25)
energy dependent digestible microbial protein: 160 FOM x 0.8 x 0.8
where FOM = fermentable organic matter nitrogen dependent digestible microbial protein: 0.9 RDP x 0.8 x 0.8
5. Questions:
What is the reason for using metabolisable or net energy systems for ruminants?
What parameters can influence the efficiency of metabolisable energy utilisation in ruminants?
How can be determined the digestible bypass protein content of feedstuffs?
How can be calculated the digestible microbial protein content of feedstuffs?

50
evaluation for companion animals
The most popular companion animals in modern-day society are the domestic dog (Canis familiaris) and cat
(Fells catus). Owning a pet has been recognised to have health and psychological benefits. Although the cat is a
true companion, the dog serves as much more than a companion. Dogs are actively used to aid blind and deaf
people as well as those confined to wheelchairs. They play a vital role in search and rescue work and are
frequently used to detect drugs and explosives. Furthermore, dogs are indispensable in the farming of sheep.
Domestication of the dog is believed to have occurred around 9000 BC while that of the cat around 3000 BC
(Rhrs 1987). Since that time many new breeds of dog and cat have appeared and currently there are
approximately one hundred and fifty different breeds of dog, while there are over a hundred different breeds of
the domestic cat. Nowadays, domestic cats and dogs are often kept in an environment where they cannot obtain
their food naturally and it is the responsibility of the owner to provide the animal with the nutrients it requires. It
becomes imperative, therefore, that the food supplied to the animal is nutritionally balanced and meets the
requirements of the animal. Failure to meet the nutritional needs of the animal for one or more essential nutrient
may result in deficiency symptoms, disease or death.
1. Anatomy of the digestive tract of cats and dogs
The dietary habit of dogs and cats throughout their evolution has resulted in morphological adaptations of the
gastrointestinal tract. The relative lengths of the parts of the intestine in the two carnivores do not differ greatly
from those in the pig. The length of the small intestine is approximately 80% of the total length of the
gastrointestinal tract with the caecum making up around 1 to 2% and the large intestine approximately 19 to
20% of the total length. The cat is different from the other three species in that it does not have a well-defined
caecum. The ratio of gastrointestinal tract length to total body length in the carnivores, however, is clearly lower
than that for the omnivorous pig.
2. Metabolic adaptations of the cat and the dog
Besides the morphological differences of the digestive tract, several metabolic adaptations have occurred in the
cat and the dog related to their dietary habits throughout evolution. This is especially true for the domestic cat,
which is believed to have evolved on a diet consisting almost entirely of animal tissues. The relatively constant
composition and the high and balanced nutrient content of such a diet is likely the reason for these metabolic
adaptations. Metabolic adaptations to a carnivorous diet are less obvious in the dog, which is consistent with the
view that this animal has evolved on a diet also containing plant material and with a less constant nutrient
content.
2.1. Protein and amino acid metabolism
There are several adaptations in the metabolism of amino acids peculiar to the cat and the dog. Besides the
normal nine amino acids, which are considered to be dietary essential for most mammals, arginine is an essential
nutrient for growing and adult cats and for dogs (NRC 1985, 1986). Consumption of a diet devoid of arginine
results in severe signs of ammonia toxicity in both species. Catabolism of amino acids generates ammonia that
cannot be converted to urea without arginine. Therefore, without arginine supplied through the diet, the
ammonia concentration in the blood increases and hyperammonaemia results.
Taurine (2-amino ethanesulphonic acid) is another amino acid regarded as an essential nutrient for the cat.
Taurine is normally derived from cysteine in the liver of mammals. However this pathway in cats is rather
limited. Recently, it was shown that dietary cysteic acid could serve as a precursor of taurine in cats. Besides the
low rate of synthesis of taurine, the obligatory use of taurine to conjugate bile acids results in a continuous loss
of taurine from the body. Taurine is not an essential nutrient for the dog as this animal can synthesise sufficient
taurine to meet its requirements.
Cats, unlike dogs, excrete several unusual sulphur-containing amino acids in their urine: felinine, isovalthine
and isobuteine. Although little is known of the latter two amino acids, rates of felinine excretion of 95 and 19
mg/day have been recorded for entire-male and entire-female cats, respectively (Hendriks et al. 1995a). These

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51
high excretion rates in entire-male cats may have a significant effect on the daily sulphur amino acid
requirement. The biological significance of felinine, or the other sulphur-containing amino acids mentioned, to
the animal is still a matter for speculation.
Another adaptation in the metabolism of amino acids is mirrored in the inordinately high dietary protein
requirement of the cat. Table 17. presents the dietary protein and total dietary essential amino acid requirements
of the growing and adult cat, dog and rat. The protein requirements for the growing and adult cat are
approximately 40 to 60% higher than those for the growing and adult dog and rat. The higher requirement for
dietary protein is not caused by a higher requirement for essential amino acids but rather a higher requirement
for the nonessential amino acids (Table 17.).
Figure 9.1. Table 17. Minimum dietary crude protein and total essential amino acid
requirements for cats, dogs and rats (Hendriks, 1996)

2.2. Essential fatty acids
Arachidonic acid is a dietary essential fatty acid for cats but not for the dog, rat or pig. Again the requirement
for this nutrient originates from the lack of activity of an enzyme in the metabolic pathway leading to the
synthesis of arachadonic acid. In the cat, the activity of the 6-desaturase enzyme is low or absent resulting in
insufficient synthesis of arachadonic acid.
2.3. Vitamin metabolism
Most mammals can convert -carotene into vitamin A. The activity of enzyme dioxygenase in the cat, however,
is negligible thereby preventing -carotene conversion into vitamin A. Cats therefore, need preformed vitamin A
in their diet to prevent deficiency symptoms.
Most mammals derive their niacin (nicotinic acid) from metabolism of the essential amino acid tryptophan. In
the cat, but not the dog, tryptophan cannot replace nicotinic acid in the diet.
How et al. (1994) showed that cats and dogs are unable to synthesize sufficient vitamin D to meet the daily
requirement. Low concentrations of 7-dehydrocholesterol in the skin of cats and dogs result in an insufficient
rate of vitamin D synthesis.
2.4. Mineral metabolism
Minerals have long been recognised to be involved in one of the most important disorders affecting the lower
part of the urinary system of cats, urolithiasis. Naturally occurring feline uroliths (urinary stones) may be
composed of magnesium ammonium phosphate hexahydrate, calcium oxalate, ammonium phosphate, calcium
phosphate, cystine and xanthine.
Strategies for preventing struvite formation in cats through dietary manipulation, either target urinary pH or the
excretion of magnesium in the urine. Lowering urinary pH to 6.0-6.4 by the addition of urinary acidifiers to
diets is one of the most effective measures to prevent the formation of struvite crystals, and has become normal
practice in commercial diet manufacture. Dry diets especially, being more prone to forming struvite in the cat,
are formulated to include urinary acidifiers such as phosphoric acid, ammonium chloride, calcium chloride and
methionine. Diets can also be formulated to contain low magnesium levels, which is the primary mineral found
in the struvite crystal.
From the information presented above on the morphology of the digestive tract and the metabolic peculiarities
of the cat and the dog, it is clear that the dog more closely resembles an omnivore than does the cat. However,
the dog has still adapted its metabolism to the carnivorous component in its diet and for this reason the dog may
be best characterised as an omnivore/carnivore. The cat on the other hand has fully adapted to a carnivorous
diet. The relatively constant composition and high nutrient content of the cat's natural diet, throughout evolution,
has resulted in several metabolic adaptations. All the nutrients that the cat cannot synthesize (taurine, arginine,

companion animals

52
arachidonic acid, vitamin A, niacin) can be found in a diet consisting of mammalian tissues. The cat, therefore,
should be regarded as a true mammalian carnivore.
3. Criteria for diet formulation
Humans have access to a wide variety of foods and will consume a number of food types to meet their nutrient
requirements. Production animals are fed compound diets formulated to support efficient growth and rarely live
out their natural lives. Cats and dogs, on the other hand and similar to humans, are expected to live a long,
happy and relatively disease-free life. However, and unlike humans, cats and dogs are often kept in an
environment whereby they cannot obtain their natural food and as a result rely on their owner to provide them
with the nutrients they need for health and well-being.
Commercial pet food manufacturers focus their efforts in diet formulation on palatability, owner's perceptions,
and providing products that offer a complete and balanced nutrition. Palatability is one of the most important
criteria in diet formulation for cats and dogs, and pet food manufacturers spend a significant proportion of their
research budget on palatability testing. Feeding their companion animal is one of the most rewarding services an
owner can perform. Seeing the enjoyment of their pet in consuming its diet gives the owner a strong sense of
fulfillment and often determines whether that particular diet (or brand) will be purchased again.
The pet owner's own perception of a diet is another important criterion in diet formulation as it also strongly
determines the re-purchasing of diets. The food an owner buys for its pet should be perceived as something
deserving of their pet. Parameters such as consistency, colour, smell, can head space and appearance of the
product are important variables in formulating diets. A dark looking paste with an unpleasant smell may be
perceived by the owner as being unnutritious food, although the diet might actually be highly palatable and
provide complete and balanced nutrition. A further measure related to owner's perception is stool quality. As
dietary ingredients have a significant effect on the dry matter content and consistency of faeces, formulation of
diets, especially for dogs, is constrained by the effects that the ingredients have on stool quality.
Marketing strategy is another important criterion in formulating diets for companion animals. Different varieties
of pet food are often produced by manufacturers, including varieties such as fish, lamb, beef and rabbit flavour.
The variety and interest concept suggests that a cat or dog needs variety in their diet.
There are positive aspects in providing a pet with a variety of diets, as it is unlikely that nutrient deficiencies
will occur on a variety of diets as opposed to the feeding of one highly palatable preferred product. For example,
vitamin E deficiency has been noted in cats due to the sole feeding of highly palatable fish or fish-based diets
(Koutinas et al. 1993). Additionally, marketing strategies influence diet formulation as commercial pet food
manufacturers often produce different brands in different price categories.
One criterion, often not as apparent as other criteria used in diet formulation for companion animals, but
extremely important, is the formulation of complete and balanced diets. Increasingly, a significant proportion of
the research budget of commercial pet food manufacturers is spent on the nutritional testing of foods.
3.1. Palatability testing
'Palatability' is a term often used in conjunction with pet foods and means 'pleasant to taste'. The importance of
palatability is evident, as a diet can provide a complete and balanced nutrition, but if it is not eaten it does not
provide nutrients to the animal and therefore is of no value. The palatability of pet foods is influenced by many
factors including food texture, food composition, ingredients, smell, taste, temperature, past experience of the
animal, heat treatment, etc.
Little information is present in the literature on palatability testing for cats and dogs. The most commonly used
palatability test in the pet food industry, the two-bowl, free choice method. In this test, animals have free access
to two bowls, each containing a different diet in amounts greater than the animal would be expected to consume
during the test period. The amount of food eaten is measured after the test period and is used to statistically test
whether there was a statistically significantly larger amount eaten from one of the two diets tested. The diet
deemed to have the highest palatability is the diet consumed in the greatest amount.
The length of the test period can be as short as two hours or as long as twenty-four hours depending on the
investigator. The number of test periods can also vary but depends on the number of animals used in a test,
available resources and the objective of the test.

companion animals

53
It is common, therefore, that commercial pet food manufacturers often have their own testing protocols, which
vary from one manufacturer to another.
After short-term palatability testing has successfully identified a preferred variety, which might be an entirely
new variety or a standard variety reformulated using more readily available or cheaper raw materials, further
preference testing can be carried out by a panel of consumers. Often an in-home placement test is used to obtain
further information on the palatability of the new or reformulated product. A typical test might involve a panel
of a hundred households of which the owners have agreed to feed a series of foods to their cats or dogs. The
owners are asked to observe how much of the diet is eaten and note other relevant parameters of interest to the
manufacturer, such as perceived colour, smell and food texture. Typically this test may run for one week. The
results of this in-home-placement test are compared with the more controlled preference tests and if the results
are suggestive of a preferred and fully accepted diet then this diet may be prepared for sale.
3.2. Tests for nutritionally complete and balanced diets: chemical
testing
Chemical analysis of pet foods and comparison with the nutrient requirements of the dog or cat should be used
in the initial assessment of the nutritional adequacy of a pet food. If, on chemical analysis, the level of a nutrient
is sub-optimal the food is deemed to provide incomplete nutrition (Morris & Rogers 1991). The nutrient
requirements of cats for growth and dogs for growth and maintenance have been reviewed by the National
Research Council (NRC 1985, 1986).
The evaluation of diets by chemical analysis, however, does not take into account the bioavailability of
nutrients. The nutrient requirements of dogs (NRC 1985) and cats (NRC 1986) are expressed in terms of
available nutrients, therefore, in most cases a diet that just meets the NRC nutrient levels would still not meet
the nutrient requirements of the animal. Calculation of the digestibility of nutrients in diets for cats and dogs
based on the digestibility of nutrients in the individual ingredients, as is practised in the livestock industry, is
possible. However, databases on by-product ingredients used in the manufacture of pet foods are limited and the
composition of these ingredients shows greater variation than is the case for similar feed ingredients for
livestock (Morris & Rogers 1991). Additionally, the often severe heat-treatment employed in the production of
commercial diets for cats and dogs may result in additional chemical damage to nutrients, and this is not taken
into account if diets are formulated based on the gross chemical composition of the prospective ingredients.
3.3. Other nutritional tests for companion animal diets
Other routine nutritional tests often carried out by pet food manufacturers are tests for urine pH and nutrient
digestibility. Manufacturers focus on the measurement of urine pH, as pH adjustment is one of the most
effective means of preventing the formation of struvite crystals.
Therapeutic diets are one class of commercially prepared diets for companion animals that are manufactured to
aid in the treatment of diseases. These diets are often sold through veterinarians and appear to be formulated on
a sound theoretical basis. Currently, diets are commercially prepared to aid in the treatment of
diseases/conditions such as chronic renal failure, obesity, feline urological syndrome, allergies, hypertension
and diarrhoea. The testing of these diets and claims made by the manufacturers, however, are not currently
subjected to regulation to prove efficacy. The testing (or lack of testing) of these diets is, therefore, at the
discretion of the manufacturer and in many countries it is possible to manufacture and market therapeutic diets
without clinical testing. Manufacturers of therapeutic diets do, however, realise the importance of clinical testing
of their diets and publish information on the efficacy of a variety of therapeutic diets for cats and dogs.
4. Prepared cat and dog foods
Dry, semi-moist and moist pet foods are manufactured from ingredients such as meats/offals (beef, fish, poultry,
deer, lamb, etc), cereal grains, meat by-products, fats/oils, vegetable protein concentrates, sugar, water,
humectants, gelling agents, emulsifiers, colourants, vitamins, and minerals. The large number of ingredients
contributes to the observed variability in the nutritional content of these types of diets. To increase shelf life,
achieve a desired physical form, and(or) increase palatability, the unprocessed mixture is either extruded, baked,
pasteurized or sterilized depending on the type of pet food that is manufactured. There is a plethora of
information concerning the influence of various heat treatments on the protein quality of feeds for pigs and.

companion animals

54
However, little information is available in the literature on the effects of the various heat treatments used in the
manufacturing of pet foods, while these are one of the most intensively processed animal foods.
Heat processing is generally believed to have a negative impact on the nutritive value of pet foods (NRC 1986).
Loss of vitamins during the production of pet foods has been extensively documented (Roche 1981). Thiamine
is particularly sensitive to heat and as a precaution compensatory amounts are added to pet foods to obtain
adequate post-processing levels of thiamine in the final product. Heat processing of canned cat foods,
furthermore, has been noted to change the taurine status of cats as moist diets increase the microbial
deconjugation of bile acids in the small intestine (Kim et al. 1996). Besides the negative impact on nutritive
value, heat processing has been noted to decrease the palatability of moist diets for cats (Heinicke 1995;
Hendriks et al. 1999).
Heat processing of pet foods can be expected to affect the protein fraction of the diet. Backus et al. (1995) found
an increase in faecal nitrogen digestibility in cats after heat-sterilisation of a canned commercial moist cat food.
This seems to indicate that heat processing would have a beneficial effect by increasing crude protein
digestibility. Determination of unmodified lysine (reactive lysine) before and after heat processing, may present
a useful in vitro method to assess heat-damage to protein. Rutherfurd & Moughan (1997) measured the
digestibility of total and reactive lysine (available) in a moist and dry cat food using a rat bio-assay. There was
no difference between the total digestible and available lysine for the dry cat food (produced by extrusion).
However, these authors found a 10% difference in the true digestibility of total and reactive lysine in a moist cat
food indicating that heat damage to lysine may have occurred.
True ileal digestibility of amino acids generally decrease with increasing heat treatment (Fig. 7.) indicating heat
damage to protein. Amino acid nitrogen and proline digestibility increased with mild heat treatment but
decreased with more severe heat processing. Aspartic acid showed the most pronounced effect decreasing
exponentially with increasing heat-treatment.
Figure 9.2. Fig. 7. True ileal digestibility of amino acids and amino acid nitrogen in
moist canned cat food after different heating periods (Hendriks et al., 1999)

It seems that dogs can digest diets more efficiently than cats. Kendall et al. (1982) determined the digestibility
of food nutrients (canned dog food, canned cat food, semi-purified, dry cat food and fresh beef mince) in both
dogs and cats. Apparent faecal digestibility coefficients for dry matter, crude protein, acid ether extract,
nitrogen-free extract and gross energy were significantly higher in the dog. Kendall et al. (1982) hypothesised
that these differences are the result of real differences in net digestive capacity between the two species. The
apparent faecal digestibility technique is an inaccurate method for the estimation of nutrient absorption in dogs.
Although the large intestine is relatively short in dogs, it appears to have a significant influence on the
determination of nutrient absorption in this species.
There are currently no data available on the influence of the large intestine on nutrient digestibility in cats.
Although the large intestine of the cat is even less developed than in the dog, published data for dogs show that
this criterion cannot be taken as evidence that dietary nutrients are not metabolized by the micro-organisms in
the large intestine.
5. Questions:
Describe the characteristics of the digestive tract of dogs and cats!
What are the main differences in the nutrient metabolism of dogs and cats?
What are the main quality criteria of dog and cat compound feeds?

55
Chapter 10. Recent advances in feed
formulation
Accurate formulation of least-cost diets for intensively raised livestock depends on knowledge of the nutrient
requirements of animals for different forms of production and on the bioavailability of these nutrients in
potential ingredients. The final manufacture of feeds for animals depends on the integration of this information
on nutrient requirements and bioavailability as well as a series of decisions that are influenced by the extent of
information available, experience of the feedstuff manufacturer and practical issues such as the ease of flow,
dustiness and palatability of the final product.
The steps involved in the manufacture of feedstuffs are as follows:
1. Establish the growth and production characteristics of the class of animal for which the feed is to be
formulated.
2. Define the tissue nutrient requirements of the animals to meet specified production goals.
3. Translate these tissue requirements into dietary specifications.
4. Define the nutrient composition and bioavailability in the range of feed ingredients potentially available for
manufacture of the feed.
5. Define the minimum and maximum inclusion limits for individual feed ingredients and nutrients.
6. Formulate the feed to meet the nominated dietary specification in a manner that will maximise profit.
7. Manufacture the feed to optimise throughput at the mill and performance by the animal.
1. Growth and production characteristics of the
livestock
The growth and production characteristics of the class of animal under consideration and the production goals
must be defined before the nutrient requirements can be specified. For growing animals, rates of deposition of
specific components of the body as well as patterns of hair or feather growth need to be identified. Factors
determining the rates of protein and energy deposition, methods for predicting the protein, fat, energy and ash
deposition in relation to body weight, food energy intake and genotype have been developed for different animal
species. Information is also available for the loss of protein in skin and in hair growth in pigs and feather growth
in poultry. This information, along with that on protein and fat deposition, has been incorporated into computer
simulation models which predict the growth characteristics and nutrient utilisation for specific genotypes and
genders of animals. For reproducing pigs, information is needed on the pattern of growth and the chemical
composition of the conceptus, including mammary tissue and uterus, as well as on the pattern of milk yield and
the composition of milk. Similarly for laying birds, information is required on the pattern of lay, egg size and
composition of the egg including its shell.
The patterns of tissue deposition in growing animals and of conceptus growth and milk yield in reproducing
animals are affected by the energy intake of the animal. For example, as the energy intake of pigs with a defined
bodyweight increases, the rate of protein deposition increases until a maximum is reached. Any further increase
in energy intake, results only in an increase in the deposition of body fat without any change in protein
deposition. Consequently, the requirement for amino acids, when expressed relative to either dietary energy or
dry matter, declines markedly when energy intake exceeds the minimum required for the maximum rate of
protein deposition. Knowledge is required, therefore, of the feed intake of the class of stock for which the diet is
to be formulated. The animals may be offered either a restricted amount of feed or be fed ad libitum. In the latter
case, information is required about how food intake capacity may be affected by the ingredients included in the
diet, and by climate, disease or the social environment of the animal. Furthermore, the rate of protein deposition
in growing animals is known to decline when they are subjected to various social stresses.

Recent advances in feed formulation

56
The rate of growth and body composition of an animal are also affected by the adequacy of available amino
acids in the diet. An amino acid deficiency will result in reduced protein synthesis, reduced growth and an
increased proportion of body fat, relative to a protein adequate diet. However, for some markets, fatter animals
may be required and the specified production goal may encourage the formulation of diets that are deficient in
protein. It is clear that many factors interact to affect the growth and production characteristics of any specific
class of stock for which a diet is to be formulated. Nevertheless, the growth characteristics and production goals
must be identified for those animals before their nutrient requirements can be specified accurately.
2. Tissue nutrient requirements to meet production
goals
Once the production goals and growth characteristics of the class of stock in question are known, definition of
the tissue nutrient requirements is relatively straightforward. The requirement of a nutrient for each body
function is simply summed. For example, the tissue requirements for protein and individual amino acids are
derived from the sum of the needs for body protein deposition, inevitable amino acid catabolism resulting in
endogenous urinary nitrogen loss, proteins and amino acids lost from the gut, amino acids needed for the
continued synthesis of skin, for hair or feather growth and, in reproducing animals, amino acids deposited in the
conceptus, mammary tissue and uterus, in eggs or secreted in milk. Traditionally for growing animals, the total
requirement for nitrogen for each body function is first established and then a fixed amino acid pattern is used to
determine the tissue requirements for individual amino acids. This approach led to the concept of ideal protein.
However, it is known that the amino acid composition of whole body protein differs substantially from that of
endogenous protein lost from the gut and of amino acids inevitably catabolised in the body.
Consequently, there is not a single amino acid pattern for ideal protein; it will vary depending on the relative
contributions of body protein deposition and endogenous losses to the total amino acid requirement. The
requirements for energy for each body function are similarly summed to determine the total tissue energy needs
of the animal. The body functions to be considered include energy deposited during body, skin and hair growth,
energy needed for maintenance, activity and thermo-regulation as well as energy deposited in the conceptus,
mammary tissue, uterus, eggs and milk. The tissue requirements for minerals and vitamins need also to be
defined. Mineral requirements depend particularly on the rate of bone growth and the need for specific minerals
for metabolic functions. Similarly, the requirements for vitamins for metabolic functions must be determined.
Traditionally, the tissue requirements of animals have been determined by a combination of the factorial and
dose response approaches, where the needs for individual nutrients for specific functions have been determined
by measuring the animal response to increasing concentrations of the nutrient in the diet, and then summed to
provide an estimate of the total requirement. The results from these 'factorial' calculations were then
incorporated into tables for use in diet formulation. However, with such an approach, it is difficult to account
adequately for the variation in nutrient requirements that results from genotype or sex of animal and for the
particular characteristics of the production environment. Consequently, computer simulation models are now
used frequently to predict tissue nutrient requirements for specific classes of animals, in defined production
systems with identified production goals.
3. Translation of tissue requirements into dietary
specifications
Translation of animal tissue requirements into dietary specifications has traditionally been one of the most
difficult tasks in diet formulation because numerous judgements must be made about the actual production
characteristics and housing environment of the animals under consideration and about the units in which the
requirements should be expressed. Judgements need to be made about the genotype and food intake of the
animals, whether their performance and, therefore, their nutrient requirements are affected by climate, activity,
disease, penning environment and other factors. In most of the earlier systems that relied on the tabulation of
dietary requirements, the effects of the animal's genetics and of environmental factors were generally ignored.
However, now with computer simulation models being used frequently to determine dietary specifications, the
interactions between the animal and its environment on dietary specifications can be readily determined.
Once the nutrient requirements for the animals in question have been determined, the second major decision is
about the units in which these requirements will be expressed. For example, amino acid requirements could be
expressed as amounts available at the tissues, truly digested, apparently digested in the small intestines or


57
apparently digested over the whole digestive tract. Once this decision is made, it is necessary to determine
whether the amino acid requirements are to be expressed as g/day, g/unit of energy or g/kg diet. Further
decisions are needed to determine whether specifications will be made for all essential amino acids and whether
they will be expressed in the selected units or as a ratio to lysine. If the amino acids are to be expressed relative
to energy, should it be gross energy (GE), digestible energy (DE), metabolisable energy (ME) or net energy
(NE)? Then, once all these decisions have been made and the units defined, a final decision is required about the
live weight range or physiological state over which the specifications should apply; that is, how many diets
should be fed to animals as they grow from weaning to slaughter; will the specifications for these diets be taken
from the beginning of the weight range, the middle or at some other point?
The most accurate dietary specifications are for amino acids available at the tissues in relation to NE. However,
the dietary specifications for the animal must be in the same units as the information on the feed ingredients that
could be considered during diet formulation. There is very little information currently on the true availability of
amino acids in feedstuffs, whereas there is a reasonable amount on the true ileal digestibility and even more on
apparent ileal digestibility. Because endogenous amino acid losses from the gut are significantly affected by
feed intake, the fibre, protein and antinutritional content of the diet and the bodyweight of the animal, the best
and most practical units for amino acids appear to be g real ileal digestible amino acid per unit of energy. NE is
a function of both the feed and the animal and is not constant for an ingredient. However, as there is little
difference in the ratio between the ME and DE contents of most ingredients and extensive information is
available for the DE content of diets for pigs and apparent metabolisable energy (AME) contents of diets for
poultry, the most practical units for specifying dietary amino acids appear to be as g real ileal digestible amino
acid/MJ DE in pigs and /MJ AME in poultry. Ideally, each essential amino acid plus the combinations of
methionine plus cystine and of phenylalanine plus tyrosine should be specified in this way. However, many feed
formulations are based on lysine expressed as g real ileal digestible lysine/MJ DE with other essential amino
acids being expressed as a ratio to lysine. Many feed formulators over specify lysine as an insurance against
deficiency and therefore over specify the other essential amino acids considered in the formulation.
Current sources of dietary specifications are tables such as ARC (1981), NRC (1988) and SCA (1987), results
from specific trials where the response of animals to increasing dietary concentrations of a nutrient has been
examined (Batterham 1993), experience of feed formulators and animal feeders (Edwards 1993) or computer
simulation models like AUSPIG (Davies et al. 1993) and the new NRC calculations (NRC 1998).
4. Nutrient composition of feed ingredients
The feed ingredients potentially available for inclusion in a diet as well as the nutrients they contain must be
specified before a least-cost diet can be formulated. Information on available ingredients and their nutrients are
traditionally contained in the databases associated with commercial feed formulation packages. This information
is frequently updated by the feed formulators. Decisions need to be made about the actual ingredients to be
incorporated in the database and which will be available for consideration during the formulation of any
particular diet. The ingredients included will vary between countries and between districts within countries
depending on the potential availability of the ingredients for particular livestock enterprises. Traditionally,
formulators in the USA have considered mainly corn and soya beans as their primary ingredients, whereas in
many European countries and Australia, a wide range of cereal grains, cereal grain by-products, legume seeds,
oil seed meals, meat and bone meals, fish meals and manufacturing by-products are included in the databases.
The number of nutrients specified for each ingredient also varies. However, it is essential that the nutrients in
the database and their units of expression be the same as those included in the dietary specifications for animal
requirements. For example, amino acids for pig diets would be expressed as g of real ileal digestible amino
acid/MJ DE to be compatible with the dimensions of the requirements. The database must include also the cost
of the ingredient delivered to the site where the feed is made and any additional manufacturing costs associated
with particular ingredients.
Although most commercial feed formulation packages have the scope to include many hundreds of nutrients,
feed formulators often consider a relatively small number, particularly when many diets are being optimised
concurrently. However, there are dangers in this practice if for instance only few amino acids are considered
during formulation of a diet and some missing amino acids could limit the production of the animals.
5. Minimum and maximum inclusion limits for
nutrients and ingredients


58
Before formulation of a diet takes place, it is important to consider whether limits should be placed on either the
minimum or maximum inclusion rates of any nutrients or ingredients. The dietary specification set the minimum
limits for most nutrients including amino acids, minerals and vitamins. However, these limits may sometimes be
altered based on previous experience of animal performance. In addition, the requirement for each mineral and
vitamin is generally not explicitly identified, but many are included in a pre-mix which will be added to the diet
at a specified rate. Sometimes specifications are made also for the minimum or maximum available energy
content of the diet.
Maximum values are sometimes set for individual nutrients, particularly minerals for which there may be
toxicities or imbalances, and for lipid and fibre because of their ability to interfere with the digestion of other
dietary components. Minimum and maximum limits are set also to ensure that the ratio of interacting nutrients
such as calcium to phosphorus or of unsaturated to saturated fatty acids is within a desired range. Furthermore,
restrictions may be imposed on the amounts of total nitrogen and phosphorus that will be included in a diet to
limit the polluting potential of the effluent.
Maximum limits are often placed on the inclusion rate of specific ingredients for reasons such as palatability,
potential for toxicity, limits to digestive capacity of the animal and potential incorporation of excessive moulds,
mycotoxins, weed seed toxins or hazardous chemical residues. Minimum limits are often imposed to force the
inclusion of ingredients that are considered to be desirable because of either their high palatability or perceived
growth promoting effects, such as milk powders and porcine plasma which stimulates antibody production.
Maximum or minimum inclusion rates of ingredients are imposed also to ensure that the final product has a
suitable physical structure to improve the ease of flow through the mill and at the farm reduce dustiness or
improve the integrity of pellets.
6. Formulation of diets to least-cost
The process of least-cost formulation of feeds using linear programming methods is relatively straightforward
and has been well defined in commercial software packages available to industry. The procedure is to match the
dietary nutrient specifications for the animal with the lowest cost supply of nutrients from potentially available
ingredients within the minimum and maximum limits set for specific ingredients. Fundamental to the process is
that the nutrient requirements for the stock and the nutrient values of the feed ingredients are in the same units.
A significant assumption in the linear programming procedure is that there is no interaction between ingredients
in the availability of nutrients. That is, the nutrient contents in all ingredients are assumed to be additive in the
formulated feed. Although this assumption is reasonable for most situations, it is not true when specific
ingredients which contain large amounts of lipid, fibre or antinutritional factors (like tannins) are included in the
feed.
A common approach to least-cost feed formulation is to minimise the cost per unit weight of feed. The difficulty
with this method is that the quantity of nutrients in a fixed weight of feed can vary widely and may not provide
nutrients at least-cost. In pig diets, DE provides an accurate measure of the total nutrient content of the diet
because amino acids and other nutrients are specified in relation to DE. The cost of diets formulated to least-
cost/tonne increased slightly as the DE content of the diet was raised from 12 to 13 MJ DE/kg and then rapidly
with further increases in the DE content of the diet to 16 MJ DE/kg. However, the cost of these diets on a
cents/MJ DE basis fell steeply from the 12 MJ DE/kg diet to a minimum at 13.5 MJ DE/kg and then rose steeply
again to 16.0 MJ DE/kg (Fig. 8.). If diets were formulated on a weight basis without a DE restriction, the 12 MJ
DE/kg diet would be chosen, but clearly at a higher cost per nutrient. However, if the diets were formulated on
an available energy basis, the 13.5 MJ DE/kg diet would be chosen. The latter method of formulation supplied
nutrients at the lower cost.
Figure 10.1. Fig. 8. Comparison the results of the cost/tonne and cost/MJ DE
formulations


59
Commercial software is now available which allows the variation in nutrient composition of ingredients to be
taken into account and to set the probability that the content of a nutrient in the formulated diet will meet the
specification. Different probabilities can be set for individual nutrients and the method is sometimes called
stochastic feed formulation. The probability constrained solutions differ from traditional linear-programme
solutions because the mean nutrient analysis, standard deviation and number of ingredients are manipulated to
provide the least-cost diet. The mean nutrient content of stochastically formulated diets is higher than that
obtained for conventionally formulated diets. Despite the theoretical advantages of being able to state the
probability that a nutrient will reach a specified concentration in the diet, stochastic feed formulation is not used
widely within the industry.
7. Manufacture of the feed
The exercise of delivering specified nutrients to the animal in question is not completed until the animal has
consumed an appropriate amount of feed, which contains the nutrients intended. If the manufactured feed either
bears little resemblance to the theoretical formulation or contains toxic and unpalatable materials, the
performance of the stock will be compromised and the effort involved in the accurate formulation of the feed
wasted.
Three issues require special attention during feed manufacture. First, the chemical composition of each
ingredient actually used in the feed mix should be known. Most commercial feed mills now use near infrared
spectroscopy to determine the moisture, protein, lipid and fibre contents of ingredients delivered and this
information is used to specify the composition of the ingredients considered in the least-cost formulation
process. In addition, ingredients should be screened for contaminants such as moulds, mycotoxins and toxic
weed seeds as well as for rancidity and other factors likely to affect the palatability of the diet, feed intake or
animal performance. Secondly, the ingredients must be accurately weighed and completely mixed. Numerous
factors such as overfilling of mixers, worn equipment, build up of ingredient residues, inappropriate mixing
times and post-mixing segregation can lead to wide variation in the uniformity of the final product.
Thirdly, the finished product should be in a form that is efficiently utilised by the animals and that flows readily
through the equipment at the mill and farm, and is not excessively dusty. It is important the physical nature of
feed on wastage (Table 18.). Pigs weighing 25 kg in groups of 10 were offered ad libitum for four weeks the
same feed either as a meal or in pelleted form. Feed intake and growth rate were unaffected by the physical form
of the feed, but feed waste was many times greater for pigs consuming the meal.
Figure 10.2. Table 18. Effect of the form of feed on the fed wastage and performance of
pigs (Hutson, 1998)

The most important factor determining the success of the feed manufacturing process is the need for vigilant and
rapid quality control capacity. It is essential that critical control point analyses are conducted on the
manufacturing operation and that standard operating procedures are documented and followed. However, the
translation of these requirements into dietary specifications still requires considerable judgement and
experience. Judgements are required about the actual factors determining the performance of stock under
commercial conditions, their likely feed intake, the units of expression of the nutrient requirements and the
actual nutrients to be included in a formulation. Considerable effort has been put into incorporating ingredients
into databases, which are normally associated with the least-cost formulation software packages. However,
judgements are also required in the setting of minimum and maximum inclusion rates for both nutrients and
ingredients. The actual formulation of the diets is straightforward, but it is important to determine whether the
feed will be optimised on a cost/tonne basis or a cost/MJ energy basis. The final manufacture of the feed
requires considerable attention to detail and quality control. Many of the steps in diet formulation have been
made more accurate and less tedious since the advent of accurate computerised prediction models.
8. Questions: 1. What kind
What kind of methods can be used for the determination of energy and protein requirements of farm animals?
What kind of characteristics of feedstuffs need to be taken into account for the accurate diet formulation?


60
What kind of factors can affect the minimal and maximal inclusion rates of feedstuffs?

61
Chapter 11. Animal nutrition and the
consumers of animal products
Animal products - meat (including fish), milk, milk products and eggs - are used principally as food for man,
although some may also be used to feed other animals. In developed countries, the by-products of the meat
industry are used to feed companion animals (cats and dogs) and may also be incorporated in the diets of farm
animals (e.g. as meat and bone meat). Milk by products are used to make milk replacers for young animals
reared away from their dams.
1. The contribution of animal products to human
requirements
After comparing the nutrient requirements of man with those of domestic animals, we must now assess the
quantities of nutrients supplied by the latter species to human diets. The contribution of animal products to
world food supplies is summarized in Table 19. In total, animal products provide about one-sixth of energy
supplies and one-third of protein supplies; meat is the major contributor, followed by milk and milk products.
The figures for individual countries and world regions differ considerably from the world averages (Table 20. ).
Thus in Europe and North America, meat consumption is 30-40 times greater than in the countries of the Indian
subcontinent, although the discrepancy for milk consumption is not as great.
Figure 11.1. Table 19. Contribution of various food groups to world food supplies
(FAO)

Figure 11.2. Table 20. Meat and milk consumption in selected countries and world
regions (kg/head/year) (FAO)

When the figures in Tables 19. and 20. are translated into nutrient intakes they show, first, that over the world as
a whole, foods derived from animals provide each person with about 1.9 MJ of energy and 28 g of protein per
day (Table 21.). These quantities are equivalent to about 16 per cent of man's energy intake and 34 per cent of
protein intake. In India, animal products supply only 7 per cent of man's energy intake and 15 per cent of
protein, whereas in the USA the corresponding figures are 28 and 64 per cent. The most important contribution
of animal products to man's requirements is that of protein. In Europe, this contribution rises from the world
average figure of 28 g/day to just over 50 g/day; in Africa, it falls to slightly less than 10 g/day. For individual
countries, the range is even greater, from 3.4 g/day in Burundi and Mozambique to 75.1 g/day in France.
Figure 11.3. Table 21. Contribution of animal products to human diets (FAO, 1998)

Animal nutrition and the consumers
of animal products

62

The main factor determining the intake of animal protein is the wealth of the human population, but this factor is
modulated by additional factors, especially the availability of alternative sources of protein and the religious
beliefs and social customs of consumers. Thus there are parts of the world, such as arctic and desert areas, where
crop production is not feasible, and the human population is largely dependent on animals for a protein supply.
In arctic areas, Eskimos eat fish and eat animals that also live on fish. In desert areas, nomadic people survive on
the products of animals such as the camel, which can live on the sparse natural vegetation.
The consumption of pig meat is prohibited by several of the major religious groups. However, sheep meat and
chicken are not commonly proscribed, except by people who choose to be vegetarians. The consumption of
milk, milk products and eggs is subject to fewer religious and social restrictions, although the extreme
vegetarians, known as vegans, exclude these from their diets. In many parts of the world fresh milk is not
consumed by adults, who then lose the ability to secrete the digestive enzyme lactase and hence to digest
lactose. They are said to be `lactose intolerant', and if they subsequently ingest foods containing lactose it is
fermented in the large intestine and causes a digestive upset.
These restrictions clearly influence global and national patterns of consumption of animal products. For
example, in India, where pig meat and beef are generally not eaten, meat consumption is very low and milk,
milk products and eggs supply a high proportion of animal protein intake. In the USA, which has a wealthy
population whose eating habits are not so much determined by religious beliefs, both meat and milk are
consumed in large amounts. Meat consumption is high also in countries with a well-developed pastoral
agriculture, such as Australia and Argentina. European countries follow close behind.
Within the worlds poorer, developing countries, there is a close relationship between social class and the
consumption of animal products, with the consumption patterns of richer people approaching those of the
developed countries. In the developed countries, however, this relationship is much less marked, as even poorer
people can afford meat and milk products. The type of meat consumed may vary between social classes,
however, with richer people eating more steaks and fewer hamburgers, hence more protein and less fat. Milk
consumption shows no systematic change with social class.
In developed countries, the patterns of consumption of animal products are liable to become ever more confused
by the growing awareness of consumers of moral objections to, and possible health risks from, such foods.
Beside vegetarians there are additional categories of people who claim to eat only a little meat, or to eat only
`white' meats (chicken and pork) and not `red' meat (beef). Others reject the white meats because they dislike
the intensive farming methods used to produce them. Concern for human health, which is discussed later,
centres on the avoidance of the saturated fats found in many animal products.
Over the world as a whole, as people and countries get richer they tend to increase their consumption of animal
products, but this eventually reaches a plateau. The level of the plateau is not necessarily the same for all: thus
meat consumption in Britain seems to have levelled off at about 70 kg per head per year, whereas the average
figure for Europe as a whole is about 80 kg, and for the USA, 115 kg. Consumer preference for animal products
may be partly based on their supposed superior nutritional value, but is probably more strongly determined by
their organoleptic characteristics (i.e. their taste and texture). Wholly vegetable diets tend to be bland and
unexciting, and meat and the other animal products are used to add variety. Improved methods of preservation
of animal products, such as refrigeration, heat processing and canning or vacuum sealing, have made it easier

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for people to enjoy a continuous supply of these products. However, the improved availability of foods in
general also means that people can experiment with exotic ingredients and culinary techniques that - like
animal products - add variety to their diet. Thus people eventually reach a point on the scale of affluence at
which they no longer need or desire to increase their consumption of animal products.
The phrase `need or desire animal products' introduces the question of their essentiality in human nutrition. Do
we really need these foods or do we just like or prefer them? The continuing successful existence of the
vegetarians, and - more particularly - the vegans, among us demonstrates the non-essentiality of animal products
for man; all the nutrients required by man can be met by foods not of animal origin. There are, however, several
major nutritional advantages in meeting man's requirements partly from animal, rather than entirely from plant,
sources. The first is that animal products supply nutrients in proportions closer to those required by man. This is
best illustrated by the essential amino acids. A growing child requires 2 g of lysine and 45 g of total protein a
day, a ratio of 4.4 g lysine per 100 g protein. For rice and wheat proteins the lysine: protein ratio is much lower
(2.8 and 3.1, respectively), so these cereals need to be balanced in the diet by a lysine-rich protein source. A
`good quality' plant protein such as that of the soya bean has a lysine : protein ratio of 6.4, but animal proteins in
milk and beef have even more favourable ratios of 8.2 and 9.1, respectively. Thus animal proteins are valuable
for supplementing the proteins of staple foods such as cereals by supplying lysine and other essential amino
acids, and this is particularly important for growing children, for whom amino acid requirements are most
critical. If lysine requirements have to be met with cereal proteins, protein intake has to be high and much of it is
wasted.
There is one essential nutrient, vitamin B12 (cyanocobalamin), which is synthesised by microorganisms and
present in animal products but virtually absent from plant-derived foods. Vegans, in particular, have to ensure
that they have a supply of this nutrient from a supplementary source such as yeast. Animal products are also
good sources of other vitamins, especially vitamin A, thiamin, riboflavin and niacin.
Another advantage of animal-derived foods for man is that their nutrients are more accessible for digestion than
those of plant-derived foods. Plant cell walls impede digestion in the stomach and small intestine and, although
they may be digested in the large intestine, the consequent release of nutrients may be too late to allow efficient
absorption. Some minerals in plant tissues are bound in compounds that resist digestion, an example being
phosphorus in phytates. Animal products are good sources of the minerals iron, calcium and zinc.
2. Ethical and environmental objections to the use of
animal products
Objections to the use of animals to feed human beings arise first from ethical considerations, a full discussion of
which is beyond the scope of this book. The primary argument, in brief, is that man has no right to exploit other
animal species. The objections to using animals are lessened if they are not killed (i.e. kept for milk or egg
production), and increased if they are kept under unnatural and perhaps harmful conditions. A second type of
ethical argument is that plant-derived foods should not be diverted to animal feeding when they could be used
directly to feed human populations that may be short of food. Until recently this argument did not apply because
farm animals were used as scavengers to convert plants and plant by-products inedible by man into human foods
(and even today, the world's crop by-products contribute about 500 Mt of dry matter per year as animal feeds).
The argument has gathered force, however, with the increasing use of cereal and other grain crops for animal
feeding. In developed countries, other than those with a predominantly pastoral agriculture, around 70 per cent
of the cereals grown are used to feed livestock, and even in developing countries (including those with food
shortages) considerable areas of land are used to grow crops for animal feeding. Over the world as a whole,
cereal usage as animal feed amounts to 115 kg per person per year; the range across countries is from 4 kg in
India and sub-Saharan Africa to 600 kg in the USA.
Objections to the use of animals to provide human food are also made on environmental grounds. Over-grazing
can destroy plant communities; demand for additional grazing can cause deforestation; the excreta of intensively
kept livestock cause pollution problems; methane from ruminants contributes to global warming. These raise
complex issues which again cannot be fully explored in this book.
The direct, nutritional objections to animal-derived foods arise mainly from two sources. First, farm animals
may harbour organisms such as pathogenic bacteria and intestinal parasites that may be transmissible to man
through the consumption of animal products. Second, some of the supposedly valuable nutrients in animal
products - fats, in particular - have been implicated in the causation of certain diseases of man.

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3. Nutrition and human health
The Merck Veterinary Manual lists 150 diseases transmissible from animals to man (known collectively as
zoonoses), but the majority of these are transferred by contact or bites, and/or are carried by wild animals. The
food-borne diseases of man that arise from farm animals form a relatively small - but nevertheless important -
group and are summarised in Table 22. The infection of man with these diseases can be minimised by various
means, the first of which involves their restriction in, or elimination from, animals. One example is the regular
use of anthelmintics to restrict intestinal parasites, and another is the slaughter of infected stock to restrict or
eradicate bovine tuberculosis. The milk-borne disease tuberculosis also provides an example of another means
of controlling infection, this being the treatment of animal products before they are consumed. The
pasteurisation (heat treatment) of milk is designed to kill tuberculosis bacilli and other bacteria. Attention to
hygiene in slaughterhouses and food stores, and appropriate cooking of meat, are also important in the control of
zoonoses.
Figure 11.4. Table 22. Some important diseases transmissible in food from farm animals
to man

Antibiotics in feeds have been used in intensive livestock systems to restrict infections, but their routine
administration is now prohibited or discouraged because of the danger of producing antibiotic-resistant
organisms.
Of the diseases listed, those regarded today as being the most important in developed countries are the enteric
infections from Campylobacter, E. coli and Salmonella organisms. Although cases of `food poisoning' have
always occurred, people today are now less tolerant of them, both mentally (food is expected to be safe) and
perhaps physically (as a generally cleaner environment has prevented the development of resistance to the
organisms responsible).
It is the possible zoonosis arising from bovine spongiform encephalopathy (BSE). The 1998 Merck Veterinary
Manual states `There is no evidence that the transmissible spongiform encephalopathies of man are acquired
from animals, but the occurrence of so-called `new variant' Creutzfeld-Jacob disease in man that has coincided
with an epidemic of `mad cow disease' in cattle and related species has inevitably cast doubt on this statement
and has led to the introduction of special control measures to prevent the transmission of BSE from cattle and to
man, and also to eradicate it from domestic animals.
The chief chemical constituents of animal products that are implicated in diseases of man are fats in general and
saturated fatty acids in particular. The diseases with which they are associated are those of the circulatory

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system that are characterised by damage to the arterial walls (atherogenesis) and the formation of blood clots
(thrombogenesis). When arteries are damaged, fibrous plaques containing lipids are formed, and these may
break away to form clots. If clots form in the blood vessels and impede the blood supply to the heart muscle
they cause what is commonly called `a coronary' (i.e. coronary heart disease); if they block the vessels
supplying the brain they cause `a stroke'; if they block the vessels of the lungs they cause a pulmonary
embolism. These conditions are frequently fatal, and if the victim survives, he or she may be severely
handicapped. Similar conditions can be caused by the rupture of damaged blood vessels.
The link between fatty deposits in the circulatory system and dietary fats is the lipid transport system that
employs lipoproteins. The lipoproteins occur in various forms, which are defined by their density and the
concentrations of these forms in blood are used to assess the risk of heart attacks and strokes. High risk factors
are high concentrations of low-density lipoproteins (LDL) and very low-density lipoproteins (VLDL).
Conversely, high concentrations of high-density lipoproteins (HDL) indicate a low risk. A high concentration of
blood cholesterol, which is a constituent of lipoproteins, is also regarded as a high-risk indicator. The
significance of these indicators is a matter of continuing research and debate. Thus some authorities consider
cholesterol level to be a poor indicator. The LDL are now thought to be dangerous when present as small
particles, a state characterised by a raised concentration of triacylglycerols.
As mentioned earlier, it is the saturated fatty acids (SFA) of foods that are regarded as the cause of a high-risk
pattern of blood lipoproteins; octadecanoic (stearic, C18), tetradecanoic (myristic, C14) and any trans acids are
considered to be the most damaging. With increasing consumption of SFA, blood levels of cholesterol and LDL
are raised. Conversely, the polyunsat-urated fatty acids (PUFA) are judged to be beneficial, although the various
`families' of PUFA differ in their effects; the omega-6 PUFA (which occur mainly in plant lipids) reduce the
blood concentration of LDL, and the omega-3 PUFA (from fish lipids) reduce VLDL. In between the SFA and
PUFA are the monounsaturated fatty acids (MUFA), such as octadecenoic (oleic, 18:1), which are regarded as
neutral or possibly beneficial to blood lipoproteins.
As the association between lipid consumption and cardiovascular disease has been exposed and explored, many
countries have produced nutritional guidelines that are intended to induce people to reduce their intake of fat and
especially of SFA. A common recommendation is that fat should provide no more than 30 per cent of total
energy intake, and that this fat should be divided equally among SFA, MUFA and PUFA (i.e. each supplying 10
per cent of energy intake). A less extreme proposal is that the ratio of PUFA to SFA (called the P : S ratio)
should be 0.5-0.8.
In Hungary, fat intake is still high, 45-50 per cent of energy intake and further changes are needed to meet the
guidelines. In the USA, fat consumption has already fallen further, to 33 per cent of energy intake, and the P :
S ratio is now 0.4-0.5. Although consumers can and do reduce their total intake of fat, they have more difficulty
in modifying the proportions in their diet of the three main types of fatty acids. Only plant lipids have the 10 : 10
: 10 ratio suggested above. The fats of terrestrial animals have a predominance of saturated fatty acids. Thus in
milk fat the ratio SFA : MUFA : PUFA is 8.5 :3.3 :0.3, and in meat, 8.3 :8.3 :2.0.
The figures given above demonstrate the difficulty - perhaps even the impossibility - of meeting the guidelines
for fat consumption with a diet containing a high proportion of animal products. The preferred strategy of those
who wish to meet the guidelines seems to be a reduction in intake of animal fat but no reduction in consumption
of the other constituents of animal products. In other words, people tend to maintain their consumption of meat
and milk (and their derivatives), but to select against the fat in these foods. Selection can be exercised by
switching from high-fat meat to that containing less fat in total and less SFA in particular; this is one reason for
the continuing replacement of beef by chicken. Fat may be trimmed from joints of meat and replaced as a
cooking aid by vegetable oils. Much of the milk consumed in liquid form has its fat content reduced to around
20 g/kg (i.e. half the 'natural' content). Of milk products, butter has to a large extent been replaced by spreads
based on vegetable oils.
Animal nutritionists, in association with animal breeders, have responded to the challenge of maintaining the
acceptability of animal-derived foods by modifying their lipid constituents. Animals are selected for leanness,
are fed to give maximal growth of muscle and are slaughtered when immature (hence having less fat). With pigs
and poultry it is possible to modify the constitution of body fats via their diet; for example, the proportions of
omega n-3 and omega n-6 PUFA can be changed. Ruminants tend to deposit saturated fat because the
unsaturated lipids of their plant diet are hydrogenated in the rumen. It is possible to introduce unsaturated fatty
acids into either their body or their milk fats by protecting dietary lipids against intervention by rumen
microorganisms, but the physical properties of the resulting fats are inevitably altered and may render them

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unacceptable to the consumer. For example, milk fat enriched with unsaturated fatty acids is more susceptible to
oxidation and hence rancidity.
The fat content of cow's milk can be reduced by feeding the cow on an extreme type of diet, typically, a diet low
in fibre, but the reduction is achieved only by upsetting the normal metabolism of the animal. This raises the
question of whether it is morally acceptable to disadvantage the animal in order to meet the perceived needs of
its consumer. Pigs selected to be ultra-lean have metabolic problems, and all pigs rely on subcutaneous fat to
provide insulation against a cold environment.
Comparisons of national populations have shown some association between the consumption of meat (especially
red meat, such as beef and lamb) and the incidence of cancer of the bowel, breast and prostate gland. However,
comparisons made within populations (e.g. between vegetarians and meat eaters) have shown no such
association, and the wholesale condemnation of meat, or other animal products, as being responsible for cancer
is therefore not justified.
The place of animal fats - and particularly the fats of ruminants - in the diets of man has been given a new
dimension by the discovery that one particular fatty acid, known popularly as conjugated linoleic acid (CLA) or
more precisely as cis-9, trans-11 octadecadienoic acid, has a beneficial role in the body. This acid has been
shown to be anti-atherogenic and anti-carcinogenic, and also to limit obesity and stimulate immune function.
The CLA is produced in the rumen as an intermediate in the bacterial hydrogenation of unsaturated fatty acids
present in the diet (hence its alternative name of rumenic acid), but it may also be synthesised in animal tissues.
It is therefore present in both milk and meat from ruminants. Ruminants given foods that contain relatively high
concentrations of unsaturated fatty acids, such as young pasture herbage, produce fats with particularly high
contents of CLA.
4. Future trends in the consumption of animal
products
Despite the arguments advanced against meat consumption - on ethical, environmental and health grounds -
world demand for all types of meat is predicted to increase steadily over the next 20 years or so. For meat in
total, consumption per person per year in the developed countries is predicted to continue to rise slowly, by 0.2
per cent per year, but for the developing countries the corresponding figure is much greater, at 1.6 per cent per
year. World demand on an absolute basis (i.e. allowing for population growth) is predicted to increase more
rapidly, by 0.6 per cent per year in developed countries and by 4.1 per cent per year in developing countries.
There are some interesting differences in the projections for individual animal species; for example, the demand
for production of pig meat in the developed countries is predicted to grow more slowly than that for poultry
meat.
It is possible that the arguments against the consumption of meat have yet to make their full impact on
consumers. However, it has been calculated that if consumers in the developed countries (i.e. those most likely
to be influenced by anti-meat arguments) were to reduce their meat consumption, world demand for meat would
still increase, by about 1.5 per cent per year. Moreover, additional projections show that, because of the adverse
economic effects on world agriculture of a reduced demand for meat in the developed countries, the partial
switch from animal to vegetable foods would not increase the world supply of food per person per day by any
significant amount.
5. Questions:
Effects of nutrition on the carcass quality of farm animals.
Differences in meat composition and its potential modification between ruminants and monogastric animals.
Which fatty acids should be increased in animal products from human nutrition point of view?
What parameters can influence the oxidative stability of animal products?

67
Chapter 12. Safety and quality of food
from animals
Quality covers many aspects such as the variety, content, composition, nutritional value, taste, freshness and
appearance of foods and recently, animal welfare has become associated with quality in the perception of
consumers. Safety issues in the 1990s include additives, veterinary drug residues, pesticide residues,
microbiological contamination and biotechnology.
1. Food charter
The increasing number of food scares and press interest fears and the way they were dealt with raised a number
of important questions about consumers' ability to have confidence in the safety of their food.
Consumers wanted to know: if livestock were free from infection, if they could be sure that the progress of food
from farm to shop was being hand led hygienically, what were animals being fed, who licensed veterinary
medicines, who inspected and set standards, who monitored these standards when they were set? In short there
were many questions of which these are only a few.
1.1. Independence
First and foremost, there must be an independent mechanism for testing, monitoring and evaluating food safety.
1.2. Safety
Consumers must be able to enjoy food without anxiety. They want the food they buy and eat to be safe. Food
safety includes such issues as additives, pesticide residues, veterinary drug residues and microbiological
contamination. The standards of safety and strategies for achieving them must be clearly set out and include a
comprehensive research programme for food safety.
1.3. Quality
Consumers expect their food to meet high quality standards. Quality covers many aspects, such as the variety,
content, nutritional value, taste, freshness and appearance of food and these points will be covered later in this
paper.
1.4. Choice
Modern consumers enjoy the benefits of a successful and innovative food industry. Consumers should continue
to have access to a wide choice of foods, but they should not be barred by low incomes from a safe and healthy
diet.
1.5. Information
Consumers must have clear, accurate, consistent and reliable information about the food if they buy. This
enables them to make informed decisions about value for money, including nutritional value. It is accepted that
consumers must take personal responsibility for what happens to the food they and their families eat after they
take it off the shelf, but to do this they must have adequate information, including instructions which enable
them to handle food safely.
1.6. Redress
Consumers need access to redress if what they buy proves to be faulty or wrongly described. They must also be
properly compensated as a matter of right if they become ill from defective food through no fault of their own.
1.7. Accountability

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68
All the agencies responsible for the enforcement and monitoring of the safety of the food supply should have the
authority and the resources to do their job effectively. The results of their surveys and findings should be
published for a wider audience. Consumers should also be told the basis on which decisions about food safety
are made.
1.8. Representation
The views of consumers should be properly represented to govemment and industry. There should be strong
consumer representation on all those committees concerned with advice, policy, priorities and strategy on
agriculture and food policy. Responsibility for achieving these principles falls on all those involved in the food
chain - the producers, the manufacturers and processors, the retailers and the distributors, the caterers and, of
course, the consumers themselves.
In the EU the prime responsibility belongs to the producer, but governments have to establish the highest
practicable standards for the food industry too. The governments must not be inhibited or dissuaded from
promptly introducing stringent regulations if the situation warrants such action. Safety must take priority over
financial considerations.
2. Representation
Safety of animal products has many strands. The animal production industry is aware of the legislation on
animal feedstuffs, about the requirements for withdrawal periods in relation to veterinary medicines, about
residue monitoring. The food industry is also aware of the moves at EU level to introduce new hygiene
standards and about the governments plan also to create a national meat hygiene service.
What we as consumers need to know is how these proposals and requirements are being monitored and
enforced. We cannot do the work. We are not the people who know how to test or inspect. So we need to have
confidence in those who can and do. We need to be sure that the best possible mechanisms are in place and that
the enforcement procedures are strong and effective.
To this end, consumers do have something to offer. We can make positive contributions to the discussions about
the standards we expect. Our views should be canvassed. We are the consumers of the products and its success
or failure depends on consumer confidence and support.
2.1. Risk
Of course, we are aware of the problems. Consumers are beginning to recognize that no product can be
absolutely safe. An element of risk cannot be eliminated. But we can cope with risk/benefit analysis if we are
presented with the facts. This is where information and labelling come in.
Full, appropriate and understandable information is required. This includes facts about any inherent risks of a
product - about unpasteurised milk, for example - so that the decision to buy and eat is a properly informed
decision. But we also need information about how to handle, store and cook foods. Many consumers still do not
know a great deal about the relationship between food poisoning and basic food hygiene.
As stated earlier, safety is now seen as an integral part of any product. It is one of the qualities we expect in
food. But what do we expect from a 'quality' food?
Quality in food - as in any product or service - has different meanings to different people. A quality product for
one shopper may be a waste of money for another. The word 'quality' on a label can mean that the food has been
produced to the highest standards of excellence or it just meets the basic legal minimum.
Recently, there have been serious efforts by consumer organizations, the food industry, the government and the
European Union to agree some definitions for 'quality' as it applies to food. In 1990, for example, the Consumer
Congress (a body which represents consumer organizations nationwide) passed a resolution calling for a
research programme on quality definitions and consumer information. Some producers have joined
certification schemes, backed by independent inspections, that set and monitor standards for the' production of
dairy, meat and fish products. In the name of quality, the EU is discussing measures to protect the names of
some traditional foods.

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So what does distinguish a quality food from the consumers' viewpoint?
There is no easy answer and the answer will vary, both from one food to another and from one shopper to
another. Consumers' perceptions of, say, a quality sausage are subjective and depend on their own personal
needs and expectations. On this basis it could be argued that a quality food product is one that meets the
consumers' requirements in every way. What are these requirements?
The Consumer Congress resolution suggested that consumers' interpretations of quality cover a very wide range
of organoleptic attributes from taste, appearance, texture and smell, through storing and cooking qualities, even
to the social policies of the food producer.
The quality of foods can be described in terms of different bands of consumer expectations as shown in Figure
9..
The inner circle - food that is gate - is the first and most basic priority. Each succeeding band represents an
increasingly sophisticated set of consumer requirements which may sometimes carry a higher price. The two
inner circles arguably represent the minimum consumer criteria for all food - not just so-called quality food.
Satisfying these basic criteria should not mean greater expense.
Figure 12.1. Fig. 9. Food quality: the consumer priorities

2.2. Safety
Consumers must be confident that the production, processing, packaging, labelling, distribution and storage of
food is within acceptable safety limits.
2.3. Nutrition
As there is increasing concern about diet- related health risks, consumers' perceptions of nutritional quality have
shifted towards lower tat, sugar and salt concentrations in food, to more fibre-rich carbohydrates and the
optimum conservation of vitamins particularly in fruit and vegetables.
2.4. Composition
Regulations that set specific compositional standards (like the minimum amount of meat in sausages or fruit in
jam) are gradually being replaced by labelling regulations, largely as a result of European Union policies. But
many consumer organizations fear there will be a levelling down and that this trend will lower food quality
standards overall.
2.5. Function
Food should be fit for its purpose. So meat sold for roasting, grilling or stewing and cream sold as whipping
cream should be suitable for its purpose. Quality here may include informative labelling to tell consumers how
to prepare and use the food.
2.6. Sensory perceptions
Many consumers decide to buy, or not to buy, on the basis of taste, smell, texture and appearance. For them, a
quality tomato feels firm, a quality avocado feels soft, fish smells fresh. Despite this people often complain
today that flavour and taste are being sacrificed for uniform size and colour. On the other hand, many retailers
assert that customers spurn unshapely or unwashed vegetables in favour of perfectly shaped, clean ones.

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These last tour attributes are closely linked but not always compatible - as recent attempts to develop tat-free
foods have shown. A fat alternative has been developed by Nutrasweet made from milk whey and egg white
proteins by a process called rnicro-particulation. The result may be of superior nutritional quality as it can be
used in a whole range of law-fat products. But it has restricted functional qualities since it cannot be deep fried
or baked. Taste trials have indicated that its sensory qualities may also be somewhat lacking.
2.7. Authenticity
This means that a load has been produced by a particular time-honoured method (such as curing, smoking,
fermenting) or in a particular place or with particular ingredients.
2.8. Convenience
Many consumers will happily pay more for products which have 'added value' such as the recipe-dish meal or
vegetables already washed and chopped, sliced or grated. This has been one of the fastest growing sectors in the
world market.
2.9. Exotic, novel, luxury
For a select group of consumers, quality means the exotic or the unusual, for which they will pay more. Vintage
champagne, truffles, and exotic, imported fruit could come into this category - although examples become
harder to find as supermarkets bring the exclusive within the reach of many. But exclusivity alone is no
guarantee of quality.
The outer circle or band in Figure 1 represents a set of ethical values which consumers can express in their
buying decisions and so exert influence over the way food is produced. Increasing concern has been shown
about the use of pesticides in crop production, intensive rearing techniques in animal production, animal
pharmaceuticals, some feeding practices and methods of slaughtering. Concern about animal welfare and the
impact of modern farming practices on the environment has provoked a growing number of consumers to
include ethical considerations in their interpretation of food quality.
2.10. Labelling and packaging
Defining the quality of food is a complex subject. But increasingly we are seeing the word quality being used on
food labels and packaging and advertisements. Some are symbols suggesting certain methods of production
(organic or free range, for example), some are stamps of approval from industry quality assurance schemes like
Food from Hungary and some are unashamedly marketing ploys.
3. Consumer awareness
Do consumers understand from the information provided what is meant grown by traditional methods or
heart friendly? It is possible that the retailers may be trying to allay consumer guilt about eating intensively
produced meat by soothing words.
There is a considerable degree of confusion about symbols denoting official schemes and others which were
purely marketing claims. The discussion group liked, for example, the symbols 'ideal for vegetarians' and
'organically grown' because they used words which seemed to make the meaning quite clear. Yet the symbol
stating that the product was organically grown had no official status and on its own did not guarantee that the
product met any recognized organic standards.
It also shows that it is not always clear to consumers when symbols are just a design feature or a claim, or when
they guarantee something about the product. Where there is any room for doubt, symbols should have words
which clearly explain the characteristics guaranteed by the symbol. It is not enough to expect that the people the
symbol is intended for will recognize it. Others may be misled if they think it has a different meaning. Symbols
should be as self-explanatory as possible.
4. Questions:
What are the main parameters that define the quality of food from the consumer point of view?

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What are the main risk categories regarding food safety?
How to inform consumers on the real quality of animal products?

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Chapter 13. Environmental impacts of
feeding monogastric animals
In different parts of Europe animal production is highly concentrated. Especially in these areas, farms have
expanded and have become more specialized. Concentration, expansion and specialization have economical
advantages; however, there are also some drawbacks. One of the main concerns is the heavy environmental load
caused by these large numbers of animals. Pig and poultry production generally is the main animal production
activity in these areas.
Environmental load can be divided into mineral load to the soil and gaseous load to the air. The mineral load is
caused by the high manure application level on the soil, caused by the unbalance between manure production
and manure requirement in these areas. Main problems arise from nitrogen, phosphorus, and heavy metals.
Surplus nitrogen leaches to ground and surface waters, causing high nitrate levels in ground water. Runoff of
especially phosphorus leads to eutrofication of surface waters. Heavy metals accumulate in the soil and will give
environmental problems in mid and long term, while residence times, depending on element and property of the
soil, can vary from hundreds to thousands of years. The gaseous load can be divided into ammonia, odour, and
methane. Uncontrolled ammonia deposition causes nitrogen enrichment of poor nature soils and acidification of
the soil, thereby affecting natural vegetation. Odour gives a problem when animal farms are located close to
residential areas. Odour is more a nuisance problem than an environmental pollutant. Methane is the most
important non-CO2 greenhouse gas. Around 20% of global methane emission is estimated to come from
ruminants and animal wastes. Methane has a high global warming potential, the impact of one molecule of
methane on global warming is 20 times that of CO2. Although nutrient losses are inevitable, nutrition seems to
be a key factor in reducing environmental pollution.
1. Nutrition and mineral excretion
The main concerning minerals are N and P. Nitrogen and phosphorus are required by pigs and poultry in a
significant amount, still most of N and P in the diet is excreted again via faeces and urine.
Nitrogen excretion can be reduced by matching the protein/amino acids content of the diet as close as possible
to the animals' requirement. Protein levels are generally higher than actually required. Safety margins in the
protein content of the diet are used to account for: 1) suboptimal amino acid ratios; 2) variations in requirement
between animals with different genotypes; 3) variations in requirement caused by differences in age or
production stadiums; 4) variations in the actual content and digestibility of essential amino acids in the diet.
Different studies show that protein content of the diet could be reduced by 30-40 g/kg without any effect on
growth rate or feed efficiency, when limiting amino acids are supplemented to the diet. Approximately 25% of
the protein in a typical corn and soybean diet can not be used, because of unbalanced amino acids. These amino
acids are broken down and the nitrogen is excreted as urea in urine.
N losses in urine also occur when energy in the diet limit protein deposition, this means that protein gain is in
the energy dependent phase. Increasing the digestibility of protein and amino acids can decrease N excretion as
well, when at the same time the protein level of the diet is reduced. In that respect it should be emphasized that
only an increase in ileal digestibility of amino acids is relevant for the update of the protein content. The
increase can be obtained by including feedstuffs with a higher digestibility, by adding enzymes or by reducing
components in the diet which causes endogenous losses. These last components are called anti-nutritional
factors and they can decrease apparent N digestibility considerably.
Protein/amino acids requirement is different for the different production stadiums. By introducing more diets for
the different stadiums a closer match can be obtained between intake and requirement. For example a three
phase feeding program can reduce N excretion by 16% when compared to a one phase feeding program for
growing-finishing pigs.
Phase feeding also can reduce P excretion, because the required concentration of P per kg of feed decreases with
increasing live weight. However, main reduction of P excretion can be obtained by increasing P digestibility. In
feedstuffs of plant origin two third of total P is present as phytic acid P, which is almost indigestible. Phytase-
supplemented feeds need little or no supplementary P. Depending on P sources and phytase dose, P-digestibility
can increase varying from 8 to 30% units.

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2. Nutrition and heavy metals
Main heavy metals of concern in the manure are copper, zinc, and cadmium. Most of the ingested Cu, Zn, and
Cd is excreted again in the manure (>90%). Already a lot has been reached to reduce Cu and Cd input on arable
land. Still, present inputs of heavy metals are too high and reducing their contents in the diet should reduce
concentrations in the manure. Other, natural, growth promoters should replace Cu and Zn in the diet. More
research about possible alternatives is necessary. By using less contaminated feedstuffs Cd in the diet can be
reduced.
3. Nutrition and ammonia emission
Ammonia in the manure originate mainly from the breakdown of urea in urine. Only a small part comes from
the breakdown of protein in the faeces. The rate at which urea is converted to ammonia depends on the urease
activity. In pure urine no urease is present, so conversion only starts when urine mixes with faeces or when it
contacts soiled floors. In a clean pen, the main part of ammonia emits from the manure pit. In soiled pens,
however, also a lot of ammonia emits from the floor. In water solutions like manure ammonia is easily ionized.
The release of ammonia from the manure is a slow process governed by factors as ammonia concentration, pH,
temperature, air velocity, and emitting surface area.
There are a few options to tackle ammonia emissions by nutritional means:
Reducing nitrogen excretion by lowering crude protein intake;
Shifting nitrogen excretion from urea in urine to protein in faeces;
Lowering the pH of manure by:
lowering the pH of faeces;
lowering the pH of urine.
3.1. Lowering crude protein intake
VThe urea excretion in the urine can be reduced by improving the nitrogen utilisation of animals. On average,
for every 10 g/kg reduction in crude protein content of the diet can result about a 10% lower ammonium content
of the manure and a 10 to 12.5% lower ammonia emission. The somewhat higher reduction of ammonia
emission compared to the reduction in ammonium content could be explained by the fact that the pH of the
manure was lowered as well when dietary crude protein level decreased. It should be remarked that water intake
can have an influence, too. At ad libitum water intake a decreased protein content of the diet reduces voluntary
water intake. This will cause the manure to be more concentrated, and therefore the difference in ammonium
concentration may be less pronounced when water is available ad libitum compared to the restricted situation.
3.2. Shifting nitrogen excretion from urea in urine to protein in
faeces
The breakdown of protein in manure is a slow process. At a manure temperature of 18 C, it takes 70 days
before 43% of protein has been broken down. In contrast the breakdown of urea to ammonia and carbon dioxide
is a fast process that covers only hours. By bacterial fermentation in the large intestine nitrogen from dietary
protein is incorporated into bacterial protein. Furthermore, urea excreted from the blood into the large intestine
can be incorporated in bacterial protein.
When increasing NSP content of the diet from 14 to 31%, a decrease in the urinary nitrogen to faecal nitrogen
ratio from 3.8 to 1.2 was found, while apparent nitrogen digestibility decreased from 85 to 75%. Combining
different studies, the relationship between NSP content of the diet and the urinaryN/faecal-N ratio was found
as shown in Fig. 10.. Enzyme supplementation (glucanase and xylanase) reduced the effect of NSP on urinary
nitrogen to faecal nitrogen ratio and pH, thereby increasing ammonia emission at the high protein level.

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74
It should be remarked that increasing the level of NSP in the diet also has negative impacts. At higher NSP
levels nutrient digestibility decreases and increases waste production, which is undesirable in animal dense
areas.
Figure 13.1. Fig. 10. Relationship between NSP content of the diet and the urine-
Ni/faecal-N ratio (Jongbloed, 2001)

3.3. Lowering the pH of faeces
With bacterial fermentation in the large intestine also volatile fatty acids (VFA) are formed. These VFA lowers
the pH of faeces and of manure. With larger proportions of NSP in the diet, not only a shift in excretion ratio
between urine and faeces, but also a lower pH of faeces and manure was found.
A decrease of manure pH with 0.12 units and a reduction of ammonia emission by 5.1% for every 100 g/d extra
intake of NSP can be achieved. Also, after a storage period of 16 days, VFA content of the manure increased
with increasing NSP content of the diet. The most profound effect on pH of manure was found when sugar beet
pulp was included in the diet of growing monogastric animals.
3.4. Lowering the pH of urine
The manure pH can also be lowered by lowering the pH of urine. The urinary pH will alter when the electrolyte
balance of the diet changes. Replacing CaCO3 by CaSO4, CaCl2, or Ca-benzoate significantly reduced the pH
of urine and manure and the ammonia emission from the manure. By replacing 6 g of calcium in the form of
CaCO3 by Ca-benzoate urinary and manure pH was lowered from 6.8 to 5.3 and from 8.0 to 6.4 respectively. In
that case, ammonia emission was reduced by almost 60%. For Ca-sulphate and Ca-chloride effects were lower
(approximately 35% reduction of ammonia emission), although the pH's of the urine were similar. Also a lower
urine pH and ammonia emission can be found by replacing di-calcium-phosphate and calcium carbonate by
phosphoric acid and calcium sulphate.
4. Nutrition and odour emission
A great number of volatile compounds have been identified in the air of pig and poultry confinement units.
Largely the same constituents have been identified in anaerobically-stored wastes. This confirms the general
assumption that malodours mainly originate from the manure. More than a hundred odourous compounds have
been identified in the air of animal houses.
Odours from animal waste can be subdivided into four chemical groups: sulphurous compounds, VFA, phenols
and indoles, and ammonia and volatile amines. Odour from manure with high total solids being dominated by
sulphides, while acetic acid and phenols predominated odours from slurry with low total solids. Sulphur
containing compounds largely contribute to the noxious odour from livestock buildings. Quantitatively VFA are
the most important groups of odour, of which acetic acid is representing approximately 60% of total VFA.
Odourous compounds are formed by the slow process of anaerobic digestion of organic substances excreted
with faeces and from the fast process of enzymatic hydrolysis of some urinary compounds. In Table 23. an
overview is given of the products that are formed by the microbial activity in manure from the main components
of urine and faeces.
Figure 13.2. Table 23. Overview of the volatile products formed by microbial activity in
manure from the main components in urine and faeces

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Dietary composition and odour production and emission have a cause-and-effect relationship and that altering
the sources and levels of crude protein and fermentable carbohydrates can be a promising approach to reduce
odour nuisance. A significant reduction in odour concentration of 77% can be found, when dietary crude protein
was reduced from 180 to 120 g/kg. Feeding a diet that better meets the protein requirement of the animals
reduces odour concentration and odour emission from the manure. A better fit to the animal's requirements can
be achieved by reducing the crude protein content of the diet and supplementing the diet with essential amino
acids. It seems that at low protein concentrations odour production does not change a lot. It might be that at low
protein concentrations protein in the large intestine becomes the limiting factor in biomass formation. This
might cause surplus of carbohydrates being converted to intermediate and odourous compounds, e.g. VFA.
There is a clear interactive effect between the crude protein content of the diet and fermentable carbohydrates on
odour concentration. It seems that an optimum balance between available fermentable protein and fermentable
carbohydrates in the large intestine of animals is a main factor in odour production. Shortage of protein or
shortage of fermentable carbohydrates will both result in formation of odourous compounds. At optimal levels,
fermentable protein is used as the nitrogen source and fermentable carbohydrates as the energy source for
biomass production.
5. Nutrition and methane emission
Methane mainly originates from bacterial fermentation of carbohydrates under anaerobic conditions. In
livestock methane mainly originate from fermentation in the rumen of ruminants. A small part is also coming
from fermentation in the colon of pigs and horses. In monogastric animals, however, a bigger part seems to
come from the manure during storage. The methane emissions measured from animal houses, expressed per kg
live weight, did not differ much between cattle and pigs.
There seems to be a close relationship between fermentable carbohydrates in the diet and methane production.
Increasing fermentable carbohydrate level in the diet to lower the pH of faeces and manure and consequently
ammonia emission, therefore will at the same time increase methane production. Although methane production
from pigs and poultry is relatively small compared to ruminants, this strategy to reduce ammonia emission
seems therefore less preferable. PH itself also influences methane production. A reduction of methane emission
with 14% was found when ileal pH was reduced with one unit by addition of acidogenic Ca and P sources to pig
diets.
6. Questions:
Methods for decreasing nitrogen excretion of monogastric animals.
Methods for decreasing phosphorous excretion of monogastric animals.

76
Chapter 14. Environmental impacts of
feeding ruminants
There is a growing awareness worldwide of the necessity to protect the environment preventing the
contamination of air with carbon dioxide, methane, ammonia, and nitrogen oxides and other gases that
contribute to the greenhouse effect. In addition, interest exists in preventing the contamination of soil and water
with excessive amounts of phosphorus (P), nitrogen (N), and potassium (K).
Like all other human activities, development and use of intensive animal production systems, including those
with ruminants, contribute to environmental pollution due to the output of waste. In intensive ruminant
production systems, energy-containing compounds produced as biomass in primary plant production are
converted to desired animal products such as meat and milk and into less desirable waste products. Waste
comprises faecal and urinary output as well as the fermentation and respiration gases carbon dioxide (CO2) and
methane (CH4). Apart from the renewable energy deposited in biomass, considerable amounts of fossil energy
are used, also resulting in a loss of CO2. In the slurry (mixture of faeces and urine), usually high amounts of
water, N, P, and K are excreted. In areas with a technologically highly developed and intensive animal
production, an efficient re-utilization of N, P, and K in primary (plant) production may become impossible by
the limited area of available land. The high water content of waste prevents its transport over large distances and
often much more N, P, and K are produced than can be taken up by primary plant production systems within the
area that can be reached economically.
Animal production is primarily driven by the input of energy, either energy in biomass or fossil energy. With
that energy varying amounts of N, P. and K are transported through the soil-plant-animal system. Transportation
through the soil-plant-animal system does not mean that N, P, and K play a passive role in the system. On the
contrary, their role is very essential, active, and often catalytic. To make or keep the soil-plant-animal system
ecologically sustainable, the three components (soil, plants, animals) of the systems should be balanced with
each other so that losses of CO2, CH4 N, P, and K are kept to a minimum. Within the animal component, the
flow of energy and its containing elements is primarily determined by nutritional strategies.
1. Environmental pollution caused by animal
production
The intensity of the nutrient flux through the animal part of the soil-plant-animal system often exceeds the
capacity of the other components of the system to efficiently utilize the elements C, N, P, and K. Besides, the
ratio between C, N, P. and K required in animal production differs from the ratio that can be efficiently handled
by the soil and plant components of the system. Apart from deposition on the soil, the imbalances also cause the
escape of C, N, P, and K to two other components essential for the system, the atmosphere and subsoil and
surface water.
The physical appearance of the four elements differs. This not only influences their availability as a nutrient for
plants and animals but is also important for the ease with which they may become dispersed in the environment.
Availability in the different compartments of the soil-plant-animal system is influenced by many different
factors and interactions.
A continuous exchange of C from solid through solute to gaseous forms and vice versa takes place through
photosynthetic, respiratory, and fermentative processes occurring in the soil simultaneously. Comparable, but
more complicated, conversions take place with N through processes known as N-fixation, ammonification,
nitrification, and denitrification. The ammonification of urea, excreted onto the soil as urine, causes ammonia to
escape into the air followed by deposition and nitrification to NO3-, which easily leaks into subsoil water
reservoirs. Alternatively, the denitrification of nitrate may cause the escape of nitrous oxides (NO,'). Ammonia
is believed to contribute to the acid deposition, and nitrous oxides are known for their harmful effects on the
ozone layer.
A special situation occurs in ruminants when C is converted into methane (CH4). Methane is also considered
harmful to the ozone layer as well as contributing to the greenhouse effect. Methane escapes to the air in
relatively small quantities, because of its much higher intensity of infrared energy absorption than CO2, and

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CH4's contribution to the greenhouse effect per gram is around 30 times higher than that of CO2. Less desired
effects of CO2 from animal production are restricted to those resulting from the use of fossil energy.
Phosphorus is an important nutrient in primary production. This element often starts its entry in the soil-plant-
animal system as an ingredient of the feed, particularly when a high concentrate to milk ratio is applied. It is
more serious for concentrate feeds if they are based on by-product ingredients. Such ingredients have a
relatively high P content, of which a relatively small proportion is absorbed and retained from it by ruminants,
and a major part is excreted in faeces. When this is subsequently used to fertilize the soil, less P is leaving the
system in animal products than enters the system with imported (concentrate) feed ingredients. In the long run
this may make P accumulate in the soil until the soil becomes saturated, after which P will filter into
groundwater or remain in surface water.
The element K is highly soluble in water. Compared with those of animals, K requirements of plants are
relatively high and again concentrate ingredients may contain high amounts, much higher than the animal can
efficiently use. Accumulation in the soil does occur only to a limited extent and excess K leaves the system in
ground and surface water.
2. Feeding Strategies and Excretion of Waste
Excretion of waste in animal production has quantitative as well as qualitative aspects. An example of
quantitative aspects is CO2 excretion, particularly that resulting from the input of fossil energy, CH4 release,
and total production of faeces and urine. More qualitative aspects are the moisture content of faeces, the ratio
between faeces and urine, the urea or total N content of the urine, the P content of the faeces, and the presence
of residues of feed and other additives. Excessive animal waste usually results from either a high stocking
density or from feeding considerably more of the nutrients N, P, and K than is required by the animals.
2.1. Energy Losses
With the animal as the basic unit, the cost of maintenance is considered an important source of waste. With
regard to energy input, a long-term feeding strategy aimed at increasing production per animal usually reduces
the input of energy from biomass per kilogram of desired product, but not necessarily the total energy input.
Higher productions need better quality feeds which often means a higher input of fossil energy. Increasing
lifetime production per animal will however reduce energy input per kilogram of product needed for
replacement. In the case of beef cattle production, energy costs of replacement can be considered as being
restricted to the costs of foetal development, which are almost negligible compared with costs of maintenance
needed to produce a carcass of approximately 400 kg in a period of up to 18 month. With milk-producing
animals, the situation is more complicated. The loss of energy per kilogram of milk produced depends on annual
production as well as on number of lactations (i.e., lifetime production) (Figure 11).
Figure 14.1. Fig. 11. The effect of annual milk yield and number of lactations on the
input of fossil energy (FE) per kilogram of milk.

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Calculations were made on the distribution of total energy input over net energy, waste from fossil energy, and
waste from energy in biomass. Assumptions were an energy input of 12.5 gigajoules (GJ) of NE for
maintenance up to 24 month of age, a deposition in the carcass (360 kg) of 9 GJ of NE, and a requirement for
milk production of 3.3 GJ/ton of FPCM (milk corrected for fat and protein content deviating from 4.0 and 3.4,
respectively). Raising the animal was assumed on the basis of an average roughage, 5.8 MJ of NE/kg of DM.
It may be concluded that with increasing milk production the output/input ratio increases, but also that part of
the increase is counterbalanced by the necessary increased input of fossil energy, which is entirely lost as CO2.
Increasing the number of lactations has a much less profound effect. Similar calculations are possible for beef
cattle. Because animal breed, growth rate, feeding intensity, and carcass composition is much more variable than
that of milk in dairy cattle, no attempt to do so was made.
Part of the energy losses are caused by the eructation of CH4. Not only is it a loss of energy, but CH4 losses also
contribute to the global warming or greenhouse effect. Methane production by ruminants has been found to be
influenced by animal size, dry matter intake, intake of (digestible) carbohydrates and other digestible dietary
components. In dairy cows, body weight, milk yield, and type of roughage influence CH4 production. The
inclusion of fat in diets for ruminants causes negative effects on CH4 losses, whereas the inclusion of
ionophores or halogenated methane analogues also causes reductions in CH4 losses.
2.2. Nitrogen Losses
Ruminal losses result mainly from an imbalance between the quantity of carbohydrates and protein degraded in
the rumen. An asynchrony between the rate at which carbohydrates and proteins are degraded may also
contribute to N losses from the rumen. Postruminal N losses occur as endogenous losses in the digestive tract
and also comprise N losses occurring in organs and tissues after absorption from the small intestine. Important
causes of endogenous losses in ruminants are protein entering the small intestine and neutral detergent fiber
(NDF) passing the small intestine. Major causes of post-absorptive losses seem to be an imbalance between the
availability of net energy and amino acids at the tissue level.
Ruminal degradation often results in a net loss of protein, due to an imbalance between protein and
carbohydrates, because of microbial synthesis. Crude protein in feed is also partly converted to CP in microbes.
Although the amino acid composition of microbial true protein is superior to that of true protein in most feeds,
this advantage is offset by the fact that 15 to 20% of microbial CP is nucleic acids, which in cattle is not
available for metabolism. Ruminal losses occur as NH3, which, after conversion into urea in the liver, is
excreted in the urine.
Dairy farming on the basis of grassland products has a relatively low N efficiency. Ruminal N losses are
particularly important under grazing conditions because cattle select for high-quality, leafy grass with a high CP
content. Besides, the intensive use of inorganic N as a fertilizer has made it possible to have high-quality;

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young, leafy forage available almost during the entire growing season. Surplus forage grown during parts of the
growing season can be conserved and fed during the winter period. The rapid degradation of CP of immature
grass is not synchronized with the degradation of carbohydrates; which creates an imbalance.
The main external input factors for N into ruminant production systems are inorganic fertilizer and concentrates.
Of these, input through N fertilizer is often more important than input through concentrates. Nitrogen
fertilization has different effects. Dry matter (DM) yield per hectare is increased in a curvilinear way, whereas
CP yield is increased linearly. The effect of N fertilization on CP content of forage is twofold: it increases the
DM yield and N content at a fixed number of days after N application and it reduces the number of days to reach
a particular stage of growth.
Forage quality can be widely manipulated by grassland management, of which N fertilization and time of
harvesting (grazing or cutting) are the most powerful tools. Other ways of manipulation are grass species or
cultivars, soil type, and season. During the second half of the growing season differences in N content in grass
due to N fertilization became much smaller, possibly because the N application in each cut was reduced. After
cutting, N content initially increases to reach a peak after about 2 wk and then rapidly decreases in the following
weeks.
Under conditions of zero-grazing outside the growing season feeding strategy depends on the application of
conserved feeds, either forages conserved by ensiling or forages and concentrates conserved by drying.
Selection can be largely excluded by feeding totally mixed rations (TMR) that are carefully balanced in such a
way that ruminal losses are eliminated and endogenous losses kept to a minimum. As an alternative, the
concentrate part of the diet can be fed frequently in small portions using a computer-controlled dispensing
system. With high-concentrate diets, scope also exists for manipulation of site of digestion as well as
influencing the fermentation pattern in the rumen. The ratio in which aminogenic, glucogenic, and ketogenic
energy is supplied can be manipulated, through which the partitioning between organs and tissues as well as the
composition of the production (milk, meat) can be influenced.
A second important site of N losses is the small intestine where endogenous protein is excreted in digestive
enzymes, bile, desquamated epithelial cells, and mucus. Causes of increased endogenous losses in ruminants are
primarily protein entering the small intestine, NDF passing the small intestine, or infection with parasites. In
sheep, approximately 10 g of extra endogenous N was passing the end of the ileum per kilogram of NDF
passing through the small intestine. Re-absorption does occur to a significant extent and varies between 50 and
75%.
A third important group of N losses is post-absorptive losses, primarily caused by an imbalance between the
availability at tissue level of (net) energy and amino acid, and to a lesser extent due to an imbalanced amino acid
profile. In cattle, liver tissue constitutes less than 5% of total body tissue, but it is responsible for 12 to 25% of
the whole-body oxygen consumption and energy expenditure and amino acids seem to be an important
substrate. Efficiency of utilization of amino acids absorbed from the blood by the mammary gland is high, and
for the majority of essential amino acids >90% of what is extracted from the blood is excreted in milk.
Feeding dairy cows according to requirements showed that the lifetime efficiency of N utilization under
standard feeding conditions remains relatively low and reaches a maximum of just below 30% at annual
productions of some 10,000 kg. Unless the animals are used for less than 3 lactations, increasing annual milk per
animal above 10,000 kg does not further increase the efficiency of protein utilization.
Increasing the number of lactations to above five, only marginally improves efficiency of utilization up to an
annual milk yield of 10,000 kg. The most effective option is to improve the ruminal and total digestibility of the
OM. This provides more substrate for microbial protein synthesis on the one hand with a simultaneous reduction
of DM passing through the small intestine, which will reduce endogenous losses on the other. It should be
realized, however, that of each gram of N captured in microbial protein, about one-third is lost either in the
faecal excretion of undigested N or in the urinary excretion of purine derivatives originating from microbial
nucleic acids. A second option is to decrease ruminal degradability of protein. Feed processing through heat
treatment, chemical treatment, extrusion, or expander cooking has been proven to be quite effective means in
that respect. Heat treatment and chemical treatment have the danger of causing an impaired intestinal digestion.
2.3. Phosphorus Losses
In ruminants, P is ingested with the feed and a varying proportion is excreted in the faeces. The proportion not
excreted in faeces is termed available P. By correcting faecal excretion for P of endogenous origin, true

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availability of P can be estimated. By far the most important endogenous source is P in saliva, which in sheep,
depending on the rumination activity, was shown to vary from one to six times the amount consumed with the
feed. Daily secretion ranges between 5 and 10 g of P in sheep and between 30 and 60 g in cattle. An amount of
60 g would provide enough P to cover maintenance and the production of 20 kg of milk. A second important
source of endogenous P is bile.
Metabolism of P in ruminants is complicated by the fact that a significant amount of P is incorporated into
microbial biomass as a component of nucleic acids and phospholipids. The ratio of N:P in microbial biomass
ranges between 6 and 7 and approaches that in milk. The proportion of feed P incorporated into microbial
biomass is not known. Recycling of P through saliva is substantial in ruminants and may exceed faecal excretion
5- to 10-fold. Although transport of P across the ruminal wall seems possible, the ruminal wall is not considered
a major site of P absorption.
In ruminants, apparent absorption of P occurs in the small intestine, and its magnitude is regulated by the
requirements of the animal. The uptake of P may be impaired by infections. Phosphorus is required for (skeletal)
growth, as a milk component (10 g/kg), and to replace P endogenously excreted in the digestive tract. Also, the
dietary Ca/P ratio may influence the efficiency of absorption. In early lactation more P is needed for excretion in
milk than is provided by the diet and the animal will mobilize body reserves, which are repleted later in
lactation. The capacity of cattle to mobilize and replete P is not well documented, but estimates suggest up to
1.5 kg, the equivalent of what is excreted in 150 kg of milk. This is considerably less than the amount of milk
that can be produced from body energy stores. Because dairy diets are largely based on grass products and
concentrates produced from by-product ingredients, dietary P levels are usually adequate, except when large
amounts of corn silage or full grains are included in the diet. Under the latter conditions it may be necessary to
supplement the diet with inorganic P.
Unlike N, the main source of P input in dairy systems is with concentrates rather than through fertilizers. A
significant part of P in feed is present as phytic acid, a form in which P is not available for monogastric farm
animals, because they lack the enzyme phytase. Although 85% of P in forages may also be present as phytate, in
the rumen it is completely hydrolyzed. For ruminants, true availability of P in forages is therefore usually high
and between 0.65 and 1.0. Virtually all P that is lost in excreta in ruminants is voided with the faeces, either in
nucleic acids or phospholipids.
Because in practice P metabolism and faecal P excretion are largely regulated by P intake, the easiest way to
reduce P loss is to reduce P input with the concentrate part of the diet. This would require the inclusion of
ingredients low in P in dairy concentrates, which normally contain between 5 and 6 g of P/kg DM. The best
option would be to include more grains in concentrates for ruminants, but these are also suitable for monogastric
animals and human consumption. Besides, the inclusion of high amounts of grain may interfere with the fibre
needs of dairy cows. Because internal inputs do not constitute any burden for the system or the environment,
home-grown feeds, such as corn silage or fodder beets, have been recommended for dairy diets. Nowadays the
development of such integrated farming systems receives much attention. Estimates suggest possible
improvements in the efficiency of utilization of N and P at the animal level from 17 to 26% for N and from 23 to
31% for P. At the farm level, improvements would be even more spectacular and efficiencies of utilization
could be raised from 15 to 39% for N and from 31 to 100% for P. Besides being internal inputs, they have the
advantage of being low in P and to some extent able to replace part of the externally purchased concentrates.
2.4. Potassium losses
Concentrates are by far the most important source of input. Input with fertilizer has been drastically reduced in
recent years, and the surplus seems to stabilize at around 80 kg/ha. Despite this, the K content in grass-based
feeds has increased by around 25% in the last few years. This is believed to be caused by the changes in manure
application methods due to legislation imposed upon farmers to reduce NH3 losses from manure. In recent
years, the application of manure has only been allowed by soil injection rather than by spreading. Application is
restricted to the growing season, which is one of the reasons K in manure is taken up by the plant very rapidly
and efficiently and accumulates in the plant. As a consequence, absorption of Mg is impaired and becomes
very low, probably below 5%, which may lead to hypomagnesemia. Incidence of hypomagnesemia has indeed
increased.
An apparent advantage of the high K levels in forages is that it dilutes urine, resulting in lower urinary N levels,
which is considered an advantage. Potassium levels of >25 g/kg DM in forage reduced urinary N concentration
by up to 30%. The dilution therefore reduces the conversion of urea into NH3, and high dietary K levels also
reduce the N concentrations in the soil where cows urinate during grazing.

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3. Implications
Animal production is inevitably associated with the production of waste and therefore some degree of
environmental pollution seems unavoidable. Various feeding strategies are available or can be developed to
minimize environmental pollution. A complicating factor is that the various forms of pollution may oppose each
other. To find optimal feeding strategies, trade-off values of the different forms of pollution have to be
determined. To reduce pollution to acceptable levels, mixed and integrated farming systems, an increased use of
home-grown feeds and in some cases extensification of animal production systems are recommended.
4. Questions:
Feeding strategies to reduce excretion of waste in ruminants.
How the methane production of ruminants can be influenced by nutrition?

82
Chapter 15. Nutritional evaluation of
the first generation genetically
modified plants (GMP)
The worldwide cultivation of genetically modified plants (GMP) increased from 1.7 million to 90 million
ha/year from 1996 to 2005 with soybean, maize, rapeseed and cotton as predominant crops. These plants are
characterized by input traits such as tolerance against pesticides or herbicides, or against insects.
Correspondingly, these plants do not have substantial changes in their composition and they are termed GMP of
the first generation.
Most of these crops are used directly or preserved in animal nutrition or as by-products from the processing
industry such as sugar beet pulp, distillers grain, extracted oil meal, etc. Therefore, nutritional and safety
assessments of feeds from GMP are one of the key questions from the public to nutritionists. Life cycle
assessments to compare the environmental impact of production and feeding of GMP in comparison with
conventional plants are of future interest.
GMP of the so-called second generation are characterized by output traits as an increased content of valuable
components (amino acids, fatty acids, vitamins, etc.), an improved availability of nutrients or a lower
concentration of undesirable substances (e.g., phytate, lignin, allergenic substances, etc.).
Recently, scientific bodies or expert panels proposed guidelines for nutritional and safety assessment of feeds
(and food) from GMP of the first generation. They gave recommendations to companies and researches for
experimental work with feed and food from GMP. Nutritional studies with feeds from GMP undertaken
worldwide.
1. Studies with Bt (Bacillus thuringiensis)-maize
Bt-maize is characterized by the introduction of a gene for a Bt-toxin which protects maize against the European
maize borer.
1.1. Beef cattle
The non-transgenic line (Cesar) and the transgenic Bt-hybrid were harvested at the wax-ripe stage and ensiled in
horizontal silos. No significant compositional differences were detected between the silages made from iso- and
transgenic lines.
For the fattening experiment, 40 male calves (German Holsteins) were raised from birth to 188 kg live weight
under standard conditions and randomly divided into two groups of 20 animals each. The performance and
slaughter results from the fattening experiment are shown in Table 24. During the fattening period, the average
daily consumption was 18.8 and 18.7 kg fresh matter from the iso- or transgenic silage, respectively. Average
daily weight gain was considered to be on a high production level.
The average carcass weight of animals of both groups was nearly identical. The leaf fat including stomach,
intestinal, channel and kidney fat was used as a criterion for the fat content in the carcass. Very high amounts of
fat were registered amounting 49.6 and 48.7 kg, respectively. However, significant differences between the bulls
of both groups were not detected.
Figure 15.1. Table 24. Fattening and slaughter results of bulls (n=20) (Aulrich et al.,
2001)

Nutritional evaluation of the first
generation genetically modified
plants (GMP)

83

1.2. Sheep
Each silage used in the fattening experiment with bulls was tested for their digestibility in four weathers. The
metabolisable energy (ME) of the silages was calculated. No significant differences concerning digestibility and
energy content were measured between both silages (Table 25).
Figure 15.2. Table 25. Digestibility coefficients and metabolisable energy content of Bt-
maize silage in sheep as compared to that of the isogenic line (Aulrich et al., 2001)

1.3. Growing and, finishing pigs
The objective of the study with Bt-maize was to determine the composition and the nutritional value of the iso-
and transgenic maize seeds fed to pigs. Both lines were analysed for proximates, starch, sugar, non-starch
polysaccharides, amino acids, fatty acids as well as for selected minerals, mycotoxins, the digestibility of
nurients and the energy content (Table 26; Table 27.).
Figure 15.3. Table 26. Chemical composition of transgenic maize seeds as compared to
that of the parental line (Reuter et al., 2002)

Compared to the parenteral line, the chemical composition, the digestibility of nutrients and the energy content
were not significantly (P>0.05) affected by the genetic modification of maize. However, the content of
mycotoxins was higher in the parenteral line. The reason for lower mycotoxin contamination in Bt-maize is the
better resistance against field infections by Fusarium spp. The lower Fusarium toxin content in Bt-maize is an
important matter of safety concern.
Figure 15.4. Table 27. Coefficient of digestibility and energetic feeding value of maize
for pigs (Reuter et al., 2002)

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84

In addition, a performance trial with 48 pigs was designed to compare the growth performance of pigs fed diets
containing either transgenic maize or its iso-line. During the growing-finishing period lasting 91 days, feed
intake and daily weight gain (Table 28.) were the same for both groups (P>0.05).
Figure 15.5. Table 28. Growth performance of growing-finishing pigs fed iso- or
transgenetic maize diets over a period of 91 days (Reuter et al., 2002)

1.4. Laying hens
For the experiments with poultry, maize cobs were manually gathered before harvesting the plants, gently dried
(40 C) and the seeds removed. The chemical composition as analysed illustrates certain differences between
iso- and transgenic maize with regard to the contents of crude protein, phosphorus and oleic acid (Table 29).
However, these differences were small, and were considered to be within the normal range of biological
variation. Six laying hens were used per group in the balance trials comprising a adaptation and a collection
period of 5 days each. The experimental diets contained 500 g maize/kg. The results of the experiment with
laying hens showed that digestibility was not significantly influenced by the maize variety (P>0.05)(Table 30.).
Laying intensity (83.5 and 83.3% at the age of 23-30 weeks) and hatchability (86.8 and 88.0% for isogenic or
Bt-maize, respectively, at 500 g maize/kg diet) were not significantly (P>0.05) influenced in a 4-generation
study with laying hens.
Figure 15.6. Table 29. Chemical composition of Bt-maize seeds and the isogenic
comparator used in the trials with poultry (Aulrich et al., 2001)

plants (GMP)

85

1.5. Broilers
Isogenic and Bt-maize were also tested in a digestibility experiment with broilers. Six birds were fed ad libitum
on diets containing 500 g maize/kg. Excreta were collected from day 30-35 of age. Protein digestibility and
energetic feeding value of the diets were also not significantly affected by the maize variety (P>0.05)(Table
31.).
Another feeding study to compare Bt-maize with the isogenic counterpart was carried out with 35 broilers over
the whole growing period of 35 days. The animals were fed a diet containing 740g iso- or transgenic maize/kg.
The results clearly showed that there were no significant differences (P>0.05) detected concerning feed intake,
digestibility, body weight gain and other performance parameters due to feeding of isogenic or Bt-maize.
Figure 15.7. Table 31. Performance of broilers fed Bt-maize or the isogenic comparator
as the principal component in the diet (Aulrich et al., 2001)

2. Bt-potatoes
Two genetically modified lines (Bt-potatoes) and a non-genetically modified control line were prepared for
analysis and feeding to broilers. A total of 27 male chicks 14-days old were randomly allotted to three dietary
treatments. Feed and water were provided ad libitum. The basal diet was formulated to contain 300 g/kg of dried
non-GM control potatoes or two lines of genetically modified potatoes (G2 or G3). The performance of chicks
was measured from day 14 to 28 of age. Summarised results of the experiment are given in Table 32. The

plants (GMP)

86
composition of the diets, feed intake, body weight and feed conversion were not significantly affected by the
potato line (P>0.05). Effects on carcass quality of broilers were also not observed.
Figure 15.8. Table 32. The influence of non-GM and GM potatoes on feed intake, final
body weight and feed conversion of male broilers (from days 14 ton28 of age) (Halle et
al., 2005)

3. Glufosinate tolerant (Pat) maize in pigs
The investigations were conducted with grains of an isogenic maize line and the corresponding transgenic
cultivar, into which a synthetically produced phosphinotricin-acetyltransferase-gene (Pat-gene) was inserted.
The nucleic acid of the codifying region was chemically synthesized. The nucleotide sequence was derived from
the amino acid sequence of the Pat-enzyme, which is produced by the bacterium Streptomyces
viridochromogenes.
Maize grain was used as test material for compositional analysis and for digestibility trials with pigs.
Compositional differences between both maize lines due to the genetic manipulation were not significant
(P>0.05, Table 33.). The levels of starch ranged from 688 to 701 g/kg DM and protein concentrations ranged
from 117 to 120g/kg DM.
Figure 15.9. Table 33. Proximate analysis, starch, sugar and NSP-composition of Pat-
maize seeds compared with those of the corresponding non-transgenic lines (g/kg DM)
(Bhme et al., 2001)

Amino acid and fatty acid profiles were determined in the three maize grain samples, to study any effects of the
genetic modification on protein and fat composition. The results are given in Tables 34. and 35. and show no
significant differences between Pat and non-transgenic grain (P>0.05).
Figure 15.10. Table 34. Amino acid analysis of Pat-maize seeds compared with the
corresponding non-transgenic controls (amino acids g/16g N) (Bhme et al., 2001)

plants (GMP)

87

Figure 15.11. Table 35. Fatty acid composition of Pat-maize grains compared with the
corresponding transgenic controls (percent of total fatty acids) (Bhme et al., 2001)

The difference technique was used in digestibility trials with pigs. Int he experimental diets 30 percent of the
DM was replaced by the different maize grains. Accross the three maize grain samples no significant differences
in digestibility of nutrients and energy content were observed (P>0.05)(Table 36.).
Figure 15.12. Table 36. Coefficient of digestibility and energy content of Pat-maize
grains for pigs as compared with those of the non-transgenic control (Bhme et al.,
2001)

plants (GMP)

88
4. Glufosinate tolerant (Pat) sugar beets
control. The genetic modifying process and the cultivation were the same as described for maize.
The sugar beet was harvested manually. The roots were washed and shredded for feeding and analysis. The tops
and leaves were chopped and ensiled in 2001 plastic silos. The resulting silages were used for the feeding
experiments after a 5-month period. The analyses for crude nutrients indicate that differences between the three
sugar-beet cultivars were detected (Table 37.). But as they are not significant (P> 0.05), they were considered
biologically not relevant. The sugar content, which contributes essentially to the nutritive value was analysed to
be the same for the non-transgenic and transgenic cultivars.
Figure 15.13. Table 37. Proximate analysis and sugar contents of Pat-sugar-beets and
Pat-sugar-beet top silage as compared to those of the corresponding non-transgenic line
(g/kg of DM) (Bhme et al., 2001)

4.1. Sheep
Digestibility experiments with sugar beet roots and top silage were carried out with wethers. The preliminary
period lasted 14 days, and faeces were collected over 10 days. The coefficient of digestibility of sugar beets was
>0.90. Significant differences between the non-transgenic and the Pat-hybrids were not detected. CF-
digestibility seemed to be improved for the Pat-hybrids by 0.033. However, a tendency towards a decline in
NFE-digestibility was found. The digestibility and the energy content of sugar-beet top silage for ruminants
showed some minor differences between the Pat-silage and controls, which proved to be statistically significant.
However, as these differences are significant only between the control and the conventionally treated
transgenic hybrid, they were considered to be biologically unimportant. This is supported by the fact that the
differences in digestibility and energy content are small.
4.2. Pigs
Sugar beets were fed in digestibility trials to five pigs of the German Landrace breed. In the experimental diets,
30 percent of the DM was replaced by the various sugar beet types. Results showed significant differences in
digestibility of the various sugar beet variants but they were minor and recorded for OM only (Table 15). OM-
digestibility was improved in the Pat-lines by 0.038 units (P<0.05), but the differences were considered to be in
the biological range. The digestibility values of crude nutrients and energy content did not show significant
differences (P>0.05; Table 38.).
Figure 15.14. Table 38. Digestibility coefficient and eneryg content of Pat-sugar beets
for pigs as compared with those of the non-transgenic control (Bhme et al., 2001)

plants (GMP)

89

5. Roundup ready (RR, Glyphosate tol.) soybeans in
pigs
The objective of the experiment was to compare transgenic (RR) full-fat soybeans with the isogenic hybrid in
growing-finishing pigs. Twelve animals were fed the isogenic hybrid and 36 pigs consumed the RR-soybeans
containing diets. The finishing period lasted from about 65 to 100 kg body weight. After slaughtering important
carcass characteristics were registered and samples from organs and tissues were taken to follow the fate of
DNA. RR-soybeans did not differ from the isogenic counterpart in all analysed constituents (Table 39.).
Feed intake, daily weight gain (836 and 859 g), feed conversion and slaughter data of pigs fed diets containing
conventional or RR soybeans were not significantly different (P>0.05) over the test period (Table 40.).
Figure 15.15. Table 39. Composition (g/kg DM) of iso- and transgenic full-fat roasted
soybeans fed to growing-finishing pigs (Flachowsky et al., 2007)

Figure 15.16. Table 40. Performance of pigs over 42 days of feeding grower-finisher
diets containing isogenic or Roundup Ready full-fat roasted soybeans (Flachowsky et
al., 2007)

6. Summary
The chemical analyses and the animal studies, which were performed with genetically modified maize, potatoes,
sugar beets and soybeans, demonstrate no significant differences as compared to the isogenic counterpart
concerning their chemical composition and their physiological production efficiency for the various species of

plants (GMP)

90
farm animals such as growing bulls, sheep, growing and finishing pigs, laying hens, broilers and growing and
laying quails. Thus, these results confirmed the substantial equivalence between feeds from transgenic plants of
the first generation and their isogenic counterpart.
7. Questions:
Describe the first generation genetically modified plants (GMP-s) used in animal nutrition!
What are the main nutritional characteristics of GMP and isogenic feedstuffs?
Results of animal experiments carried out with GMP and isogenic feedstuffs.

91
Chapter 16. Nutritional evaluation of
the second generation genetically
modified plants (GMP)
GMP-s of the second generation are characterized by:
Increased contents of desirable substances (e.g., amino acids, vitamins, fatty acids, minerals, enzymes).
Decreased contents of undesirable substances (e.g., mycotoxins, alkaloids, glucosinolates, lignin, phytate).
At present, detailed standardized test procedures are not available to investigate feeds from the GMP of the
second generation. Depending on the claim of changes due to the genetic modification, the experimental designs
must be arranged to demonstrate the effects. Different experimental designs are necessary to demonstrate the
efficiency of changed nutrient constituents:
Bioavailability or conversion of nutrient precursors into nutrients (e.g., (3-carotene).
Digestibility/bioavailability of components (e.g., amino acids, fatty acids, vitamins).
Efficiency of substances which may improve digestibility/availability (e.g., enzymes).
Utilization of substances with surplus effects (e.g., prebiotics).
Improvement of sensory properties/palatability of feed (e.g., essential oils, aromas).
However, the genetic modification might not only increase the content of intended desirable substances. There
are indications that side effects may occur and cause unfavourable effects. Such secondary changes should be
considered in the nutritional and safety assessment of GMP of the second generation. Specific animal studies as
the basis for comparative approaches seem to be necessary to deal with these questions.
1. Increased myristic and palmitic acid in rapeseed for
pigs
Rapeseed with modifications in the fatty acid pattern was analysed for its composition and its feeding value for
growing-finishing pigs in comparison with the non-modified counterpart. The objective of this genetical
modification was to produce rapeseed for technical purposes. But as the by-product is intended to be used as
feedstuff, the seeds were analysed for composition and energetic feeding value for pigs.
Except for the fatty acids, the GM-rapeseed showed only marginal differences in nutritional composition, but
the glucosinolate (GSL) content was increased (Table 41.). The digestibility and the energy content of the diets
containing 150 g iso- or transgenic rapeseed/kg fed to five growing and finishing pigs each remained unaffected
(Table 41).
Figure 16.1. Table 41. Chemical composition of iso- and transgenetic rapeseed

Nutritional evaluation of the second
plants (GMP)

92

The higher concentration of myristic and palmitic acid of the transgenic rapeseed fed to pigs had a negative
influence on feed and energy intake and consequently daily weight gain (Table 42).
Figure 16.2. Table 42. Coefficient of digestibility and energy content of rapeseed-based
diets and performance parameters of pigs (n=10, 32-105 kg body weight)

The reason for this depression was due to the fact that the genetic modification was associated with higher
concentrations of undesirable substances. These results are an excellent example that genetically modified plants
with output traits need a complete compositional and nutritional assessment. This is also supported by results
obtained from GM-potatoes.

plants (GMP)

93
2. Inulin synthesizing potatoes in pigs
The ability to synthesize high molecular weight fructan as inulin was transferred to potato plants. As the fructan
pattern of tubers from this transgenic potato plant represents the full inulin spectrum of artichoke roots, the
tubers were intended to be used as a prebiotic functional food in human nutrition. The inulin concentration in the
dry matter of the transgenic tubers amounted to 50 g/kg.
Proximate composition, minerals and amino acids did not show significant differences between lines (Table 43).
However, the starch content decreased as inulin was stored, indicating that the storage-capacity of carbohydrates
was not affected by genetic modification. The total alkaloid content of the transgenic tubers was about 25%
higher than that of the isogenic potatoes. In agreement with the data presented for rapeseed, the results confirm
that substantial genetic modifications might be associated with altered concentrations of undesirable substances,
and therefore increased attention should be paid to this fact and in additional safety studies.
Figure 16.3. Table 43. Selected proximate analysis, starch, macro-elements, amino acids
and glycoalkaloids of transgenic inulin synthesising potatoes compared with those of the
parenteral line

Digestibility depressions of some nutrients of the inulin-synthesising potatoes were detected and
correspondingly a lower energetic feeding value was measured (isogenic: 14.60; transgenic: 14.34 MJ ME/kg
DM, P>0.05). The lower production potential of the silage from transgenic potatoes was also confirmed in the
feeding test. The average daily liveweight gain of the pigs fed transgenic silage was 43 g lower as compared to
controls (P>0.05). The results show the reduced energy and prebiotic potential of the GM-potatoes, but they are
not significantly different from those of the control (P>0.05).
3. Fate of DNA
The fate of DNA especially transgenic DNA during feed processing and in the animal received attention after
studies by Schubbert et al. (1994, 1997, 1998) who found that DNAfragments after feeding of phage DNA to
mice were absorbed and detected in blood, liver, spleen and other organs and tissues. This makes it necessary to
investigate the fate of plant DNA in farm animals, especially as far as recombinant feed plants are of concern.
Beginning with the first experiments the fate of DNA in the animal body was studied (Einspanier et al., 2001).
Meanwhile the fate of DNA was studied in dairy cows, beef cattle, growing-finishing pigs, laying hens, broilers
and quails fed Bt-maize, Bt-potatoes, RR-soybeans, or inulin potatoes.
The most important results are summarized in Table 44. In agreement with other findings, most DNA is
degraded in the gastrointestinal tract, but some fragments have been found in animal tissues. However, no

plants (GMP)

94
residues of recombinant DNA were detected in any organ or tissue samples including eggs and milk obtained
from animals fed with feeds from GMP (Table 44). However, in the case that plant DNA-fragments should be
absorbed, it might be that transgenic DNA-fragments are also absorbed. If this occurs, the frequency is likely to
be extremely low.
4. Questions:
Describe the second generation genetically modified plants (GMP-s) used in animal nutrition!
What is your opinion on using GMP feedstuffs in animal nutrition?
Figure 16.4. Table 44. Studies of the transfer of foreign DNA fragments into farm
animals

95
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