Module 4

The Botany of Wheat: anatomy, growth and development and physiology

I. Introduction II. Zadoks Decimal Growth Stages a. Germination b. Seedling growth c. Tillering d. Stem Elongation e. Booting f. Ear Emergence g. Flowering h. Milk Development i. Dough Development j. Ripening III. Roots IV. Vernalization and Photoperiod V. Physiological Processes Driving Growth and Development

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I. Introduction.

In seed time learn, in harvest teach, in winter enjoy.

William Blake

Whatever kind of seed is sown in a field, prepared in due season, a plant of that same kind, marked with the peculiar qualities of the seed, springs up in it.
Guru Nanak The anatomy of and the growth and development of wheat has been studied extensively and will be presented here in review. Understanding of how the wheat plant grows and develops along with the capacity to utilize the terminology that accurately describes the plant and its life cycle is important. Firstly, an acquaintance with these subjects aid in communication with others that may be cooperating to some extent with the breeding objectives. Secondly, proficiency in these topics will enable the breeder to identify traits that may be associated with improved performance. Physiological criteria are commonly used in breeding programs, a widespread example being the selection for semi-dwarf cultivars. Selection for reduced height has improved lodging resistance, and increased the harvest index (HI) of wheat by increasing the partitioning of biomass to the ear and developing grain. The physiological bases behind superior performance are just beginning to be uncovered. A number of these traits have a strong association to performance and have high heritability and can therefore be used in the selection process. These physiological traits will be discussed in the modules: Breeding for Increased Yield Potential, and Breeding for Drought Stress. An understanding of the wheat plants growth and development will aid the breeder in selection for disease resistance. Different diseases will attack the wheat plant at different developmental stages, and knowing the proper time to rate disease is paramount in applying accurate selection pressure. Selection for wheat plant development can also aid in developing a cultivar that can better compete with weeds by shading them out more quickly, or avoid seasonal environmental stress by earliness of maturity. In short: an understanding of the wheat plants anatomy, growth and development and physiology can give the breeder a wider range of tools to aid in the science of breeding, and a better intuition to aid in the art of plant breeding.

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II. Zodaks Decimal Growth Stages. There are several scales or developmental codes for wheat that describe visible growth stages. Haunss scale (Haun, 1973) is useful in defining vegetative growth stages, and Feekes scale (Large, 1954) provides a good description for both vegetative and reproductive stages. However, Zadoks scale is the most comprehensive and easiest to use. It describes all stages of the cereal growth cycle, incorporating characteristics not considered in other scales. Growth is a complex process with different organs developing, growing and dying in overlapping sequences and it is easier to think of it as a series of growth stages as in the Zadoks scale. This has 10 main growth stages, labeled 0 to 9, which describe the crop; and each main growth stage can be further described using a second digit, labeled 0 to 9 (Table 1). After emergence, all developmental stages are based on observations on the main shoot. After stage 40 the stages of the main shoot and tillers become similar, and the stages are determined by viewing the whole plant. Stages 70 to 93 are determined by the development stage of individual kernels or grain in the middle of average spikes. Although Zadoks growth stages parallel growth and development along time they do not necessarily follow a sequence per se as decimal stages 1 and 2 as well as 2 and 3 occur in parallel, and a plant in a single moment could be described by two or three stages (see figure 2) II. a) Germination: Zodaks 0 Imbibition describes the process of the seed taking up moisture from the soil in order to break dormancy and begin germination. The minimum water content required in the grain for wheat germination is 35 to 45 percent by weight (Evans, 1975 WGP). Germination may occur between 4 and 37C (optimal germination temperatures range from 12 to 25C). Upon hydration the scutellum (see figure 1) begins to mobilize its own starchy reserves along with secreting enzymes that break down the starchy endosperm. The digested endosperm is absorbed by the scutellum, and nutrients are conveyed to the growing embryo. The radicle will first emerge and begin to grow downwards along with about four other seminal roots. The coleoptile emerges shortly after the radicle. The coleoptile forms a sheathing structure through which developing leaves grow. The coleoptile increases in length until it emerges through the soil surface, where it ceases to grow. On average the coleoptile reaches 5cm in length. Selection at this stage will most likely be by natural means as those genotypes that do not contain suitable vigor will either not germinate or will not reach the soil surface. CIMMYT places selection pressure at this stage for drought tolerance by deep planting. Soil moisture, in most cases, will be greater at depth; and deep planting is a way to expose seeds to moisture that may not be available closer to the surface (a possible problem is water-stressed environments). II. b) Seedling Growth: Zodaks 1 Zodaks 1 describes leaf emergence. Leaf primordium appears first as a bump on the flank of the shoot apex. The leaf primordia grow laterally and acropetally becoming a cowl shaped sheath. The sheath-like leaf grows upward as a conical cap over the shoot apex and younger leaves develop within in the same fashion. When the leaf is about 20 mm long, the ligule develops separating the leaf blade (lamina) from the sheath which

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Figure 1

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will remain wrapped around the stem. The ligule is a thin membranious outgrowth generally regarded as a protective structure which prevents rain, dust, insects and other foreign material from entering and accumulating within the sheath; protecting younger, underdeveloped tissues from damage. Associated with the ligule and located in the same position are the auricles which partly wrap around the stem (fig. 2). Leaf shape and size change with leaf position. The first leaf on the main shoot has parallel sides to within 1 cm of the tip making it relatively blunt ended. The leaves above the first have more or less parallel sides for about two-thirds of their length above which they taper to a sharp point. The last leaf produced upon the culm, the flag leaf, tapers from about the lower third, giving the leaf an elongated ovate shape. As the life cycle of wheat progresses, lower plant leaves die due to shading, drought, disease, or normal maturity. The flag leaf is the last to remain green and accounts for 80 percent of the carbon dioxide assimilation of grains. Figure 3 demonstrates Zadoks notation in relation to leaf development. Studies on historical sets of cultivars suggest that leaf shape and orientation have contributed to genetic progress in yield and this will be discussed in the module: Breeding for Increased Yield Potential and Yield Stability. Also certain metabolic processes that occur in the leaf will be discussed later along with their potential use as indirect selection criteria for yield and drought tolerance will be covered in the modules: Breeding for Increased Yield Potential and Yield Stability, and Breeding for Drought Tolerance.
Fig. 2 Vegetative Structures Of Wheat

Flower Spike Hollow Stem Culm Node Internode Stem (culm) Leaf Blade Ligule Auricle Sheath

Sheath Tiller Stem section and leaf sheath

Figure 3 Zodaks 1

FAO Irrigated Wheat. Howard M Rawson and Helena Gmez Macpherson

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Table 1 - Decimal code used to quantify the growth stages in cereals

Code 0 0.0 0.1 0.2 0.3 0.4 0.5 1.0 10 11 12 13 14 15 16 17 18 19 2.0 20 21 22 23 24 25 26 27 28 29 3.0 30 31 32 33 34 35 36 37 Description Germination Dry seed Start of imbibition Imbibition complete Radicle emerged from seed Coleoptile emerged from seed Leaf just at coleoptile tip Seedling growth First leaf through coleoptile 1 leaf unfolded 2 leaves unfolded 3 leaves unfolded 4 leaves unfolded 5 leaves unfolded 6 leaves unfolded 7 leaves unfolded 8 leaves unfolded 9 leaves or more unfolded Tillering Main shoot only Main shoot and 1 tiller Main shoot and 2 tillers Main shoot and 3 tillers Main shoot and 4 tillers Main shoot and 5 tillers Main shoot and 6 tillers Main shoot and 7 tillers Main shoot and 8 tillers Main shoot and 9 or more tillers Stem elongation Pseudo-stem erection 1 node detectable 2
nd rd st a

Code 38 39 4.0 41 43 45 47 49 5.0 51 53 55 57 59 6.0 61 65 69 7.0 71 73 75 77 8.0 83 85 87 9.0 91 92 93 94 95 96 97 98 99

Description Flag leaf ligule just visible Flag leaf ligule just visible Booting Flag leaf sheath extending Boots just visible and swollen Boots swollen Flag leaf sheath opening First awns visible Ear emergence First spikelet of ear just visible One-fourth of ear visible One-half of ear emerged Three-fourths of ear emerged Emergence of ear complete Flowering Beginning of flowering Flowering half-way complete Flowering complete Milk development Seed water ripe Early milk Late milk Dough development Early dough (fingernail impression not held) Soft dough Ripening Seed hard (difficult to divide with thumbnail) Seed hard (cannot dent with thumbnail) Seed loosening in daytime Seed over-ripe; straw dead and collapsing Seed dormant Viable seed giving 50% germination Seed not dormant Secondary dormancy induced Secondary dormancy lost
c b

Medium milk

Hard dough

node detectable

3 node detectable 4* node detectable 5th node detectable 6 node detectable Flag leaf just visible
th

(a) Winter cereals only. (b)An increase in the solids of the liquid endosperm is notable when crushing the seed between fingers. (c) Fingernail impression held; head loosing chlorophyll. Source: Zadoks et al., 1974.

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II c) Tillering: Zadoks 2 The wheat plant has the ability to tiller (produce lateral branches). The number of tillers that will be generated depend on a number of factors including: genetic, population density, sowing date, and the availability of water and nutrients. Tillering normally starts when leaf 3 is fully expanded and leaf 4 is emerging on the main shoot. The main shoot bears primary tillers in the axils of its leaves and can be described in relation to the leaf number (TC emerging from the axil of the coleoptile, T1 emerges from the axil of leaf 1, T2 emerges from the axil of leaf 2 etc.). Each primary tiller has the potential to bear a number of secondary tillers and can be described in reference to the leaf number of the primary tiller (T11 is the tiller borne in the axil of leaf 1 of tiller 1). Of the tillers that emerge, only a proportion will survive to produce seed, the rest dying without producing an ear, possibly due to competition for resources. Generally about eight tiller buds will form, but only three or four will develop into full size tillers that produce seed. Tiller appearance generally ends just before stem elongation begins. Figure 4 diagrams the nomenclature used for describing tillers along with drawings of plants typical of Zadoks stage 2. Tillering is under genetic control and varies among cultivars. Selection based on number of fertile tillers may be the source of genetic gains for different cropping systems (basin or raised bed). Recent studies also show that a tiller inhibition gene (tin) may prove useful in regions that are regularly subjected to terminal drought (Duggan 2005a, Duggan 2005b) and will be discussed in the module breeding for drought resistance.
Fig. 4 Tillering a) Nomenclature for leaves and tillers. b) drawing of wheat plant at Zadocs stages 1 and 2

Source: Kirby and Appleyard, 1985. (Courtesy of Kluwer Academic Publishers)

FAO Irrigated Wheat. Howard M Rawson and Helena Gmez Macperson

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II d) Stem Elongation: Zadoks 3 Stem elongation usually begins between the late double ridge and terminal spikelet stage (discussed below). The elongation of the stem coincides with the growth of leaves, tillers, roots, and the inflorescence, which raises questions about possible competition for the assimilate supplies. Lower internodes of the stem remain short. A spring wheat with 9 leaves will begin elongation with the 4th internode, while a winter wheat that has more leaves will begin elongation at a higher numbered internode. When an internode has reached about half its length the internode above will begin to elongate. Each succeeding stem internode is progressively longer and the sequence of elongation will continue until anthesis. The peduncle, topped by the ear, is the final stem segment to elongate and can account for as much as half of the total stem length. The height of wheat ranges from 30 to 150 cm and is influenced by genotype and environment. Reduced height genes (Rht) affect internode length and have had a larger impact on modern wheat production than any other physiological trait to date. Until the time of stem elongation the shoot apical meristem had been the initiation site for vegetative growth. Just before stem elongation begins the shoot apex begins to initiate spikelet primordial which marks the commencement of reproductive growth for the wheat plant. Each of the primordial initiated on what will become the ear has two parts (the double ridge, see fig 5). The lower, smaller ridge is a leaf primordia, the further development of which is more or less completely suppressed. The upper larger ridge eventually differentiates to become the spikelet. The double ridge stage occurs when from 40 to 80 percent of the spikelets have been initiated. After about 20 to 30 spikelet primordia have been initiated, the final number of spikelets is determined by the formation of a terminal spikelet. Each spikelet has from 8 to 12 floret primordia in the central part of the spike, while basal and distal spikelets have from 6 to 8 florets.
Fig. 5 - Successive stages of shoot apex development from a vegetative apex (a) to double ridge (c) to terminal spikelet stage (f)

b
dome leaf primordia

Site of spikelet ridge Lower leaf ridge

c
Axillary spikelet ridge Lower leaf ridge

leaf primordium Terminal Spikelet Lemma, Floret Lower glume

e d
Spikelet meristem Glume Primordia Spikelet meristem Floret Lemma Glume

f
Spikelet meristem Lemma, Floret 3 Stamen lemma
Source: Adapted from Kirby and Appleyard, 1987. (Courtesy of Arable Unit RASE)

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Spikelet development on the microscopic head is usually completed by the time the first node is 1 cm above the soil surface. The terminal spikelet is produced at about Zadoks stage 31. A rapid loss of younger, poorly developed tillers also normally starts at this stage. The stem elongation or jointing stage comes to an end with the appearance of the last (flag) leaf. II e) Booting: Zadoks 4 The developing head within the sheath of the flag leaf becomes visibly enlarged during the booting stage (figure 5). The booting stage ends when the first awns emerge from the flag leaf sheath and the head starts to force the sheath open. Spike growth occurs after the terminal spikelet is formed and stem elongation has begun. Spike growth is slow in its early stage and increases greatly about the time the ligule of the flag leaf becomes visible (Krumm et al., 1990 FAOBW44). This time of rapid spike growth is important in determining yield at harvest as an environmental stress can decrease the supply of assimilates and contribute to floret death (see figure 6). Floret abortion, which starts in the boot stage and finishes at anthesis, occurs when stem and peduncle are at maximum growth rate (Siddique et al., 1989FAOBW44). Meiosis in wheat, which originates the pollen in the anthers and the embryo sac in the carpel, coincides with the boot stage. This stage is very sensitive to environmental stresses. In wheat meiosis starts in the middle of the spike, continuing later above and below this zone (Zadoks et al., 1974 FAOBW 44).

Fig. 5 Wheat inflorescence at Zadoks stages: 4 Booting, 5 Ear Emergence, and 6 Flowering.

FAO Irrigated Wheat. Howard M Rawson and Helena Gomez Macpherson

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Fig. 6 External and Internal Stages and When Yield is Formed

Figure From: FAO Irrigated Wheat. Howard M Rawson and Helena Gmez Macpherson

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Possibilities for selection of a trait to increase performance are related to the phenology surrounding spike growth. Phenology refers to the study of cyclic events in nature and in this case it is addressing the length of time for spike development. This will be further discussed in the module: Breeding for Yield and Yield Potential. The duration of spike growth has been shown to be associated with fertility and is under genetic control which varies among cultivars (Miralles and Slafer 2007). II f) Ear Emergence: Zadoks 5 As the stem continues to elongate, the still growing spike begins to emerge from the flag leaf sheath (figure 5 heading). The time required for the spike to completely emerge from the sheath is highly dependent on environmental conditions and can range from one to five days. Throughout the preheading period, differences in the duration of the various developmental phases among shoots on the same plant help synchronize development. This means a difference of several weeks between emergence of the main shoot and a tiller is reduced to a difference of only a few days by the time the heads emerge from the flag leaf sheaths. At this stage final developmental events take place in the ovary, and pollen mother cells leading to anthesis. II g) Flowering: Zadoks 6 The wheat inflorescence is a spike bearing spikelets at the nodes (see figure 7). There is one spikelet per rachis node. Each spikelet is surrounded by a pair of glumes and contains two to six florets. Found in each floret, enclosed by the palea and the lemma, are three anthers, the stigma, and the ovary. Anthesis, or the shedding of pollen occurs about three to ten days after the ear emerges from the flag leaf sheath (see figure 5 flowering). The lodicules (see figure 7) swell causing the palea and the lemma to open. The stamen filaments elongate and the anthers dehisce, or shed pollen. The whole process is complete within about five minutes (Percival, 1921 FAOBW 34). Anthesis begins in the early morning and continues throughout the day. For a single spike this process takes four to seven days. Anthesis begins in floret one of the spikelets in the upper two thirds of the spike. The following day anthesis progresses to the first floret of the basal spikelet and to the second floret of the upper spikelets. The progression continues ending with anthesis occurring in the last fertile floret of the basal spikelets (Evans et al., 1972FAOBW34). Within a single plant, anthesis occurs first in the main shoot with the anthesis of tiller spikes commencing within three or four days. Within ten days or less, depending on environmental conditions, a single wheat plant completes anthesis. Cultivars differ in the degree to which the lemma and palea are separated. Most often, individual florets are self pollinated. However, while the flower is open, foreign pollen may enter, resulting in about one or two percent cross-pollination. Lodicules lose their turgor and the florets close within an hour of opening. The lodicules degenerate after the first opening, but the ovary will swell and the floret may open again. The stigma of an unfertilized floret will remain receptive for up to five days after the time of anthesis (emasculated or male sterile floret).

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Fig. 7 A wheat Spikelet

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II h) Milk Development: Zadoks 7 The pollen grain contains two nuclei, one generative and one vegetative. With successful pollination, the vegetative nuclei will begin developing a pollen tube which will create a path through the style to the ovule and entering the embryo sac through the micropyle. The generative cell follows the path created by the tube cell and along the way replicates and divides to produce two sperm cells. One sperm cell unites with the egg. This union produces a zygote which will become the embryo. The second sperm cell will unite with the central cell which will become the 3n endosperm which will eventually be the source of wheat flour (figure 8a). The seed increases in size about three fold in the four days after fertilization as the tissues surrounding the embryo swell. The growth is caused by an expansion of the cells rather than cell multiplication. In these first few days development of the zygote is slow, cell division occurs producing a globular shaped embryo (figure 8b). Also in the first four days, the 3n nuclei replicate and divide fueled by the antipodal cells. This phase of free nuclear division will, in part, determine the final number of cells in the endosperm. The time between four and ten days after fertilization is called water-ripe and is described by Zadoks stage 71. At this time the inside of the grain has little structure and when opened appears to contain only water. At this stage the nuclei continue to divide rapidly and cell walls begin to form. By 7 days after fertilization the embryo begins to show signs of differentiation. Eleven days after fertilization marks the medium milk stage, Zadoks 75, and the first stage of grain filling. By 16 days after fertilization lipid and protein bodies can be found in the endosperm as well as A-type starch grains. The cell layers that surround the embryo sac continue to change. The cell walls thicken and the aleurone becomes recognizable. The embryo is developing quickly and takes on an elongated shape. At 16 days after fertilization the scutellum is clearly defined and the embryo will begin to use the endosperm starch reserves for its development. II i) Dough Development: Zadoks 8 At 21 days after fertilization the outside of the grain begins to turn from green to yellow and marks the soft dough stage, Zadoks 85. Cell division of the endosperm has ceased and A-and B-type starch granules that formed during the medium milk stage are packed into cellular compartments. The inside of the grain is still moist but the contents are now semi-solid. The embryo, continuing to feed off of close-by endosperm, has grown to half its size. Twenty-one to thirty days after fertilization the caryopsis enters the hard-dough stage, Zadoks 87. The grain takes on a golden color as protein and starch accumulation ceases followed by the death of the endosperm cells. The embryo is now fully developed, but will continue to receive storage reserves until the grain begins to dry and a state of dormancy in initiated. II j) Ripening: Zadoks 9 Ripening or dry down occurs between days 30 and 40 after fertilization. The water content of the grain drops at a continuous rate, but there is a wide degree of variability of the time for complete dry down between different grains in the ear and different ears in the crop. The moisture of the grain must be watched closely in order to

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harvest at the optimum time. Further air drying is often used after harvest to better control the final moisture content of the grain. If the process of desiccation is not controlled properly, the grain may begin germination destroying the quality of the grain. Pre-harvest sprouting is dependent on both environmental and genetic factors and is a major problem that occurs most often in white wheat exposed to moisture at harvest time. Grain moisture at harvest is generally 20% and must be further dried to about 14% for proper storage.

III. Roots
So far described have been the aerial parts of the wheat plant, and a few words must be said about the development of the root system. The root system of wheat is, of course, important for its performance in all settings. It is of critical importance in developing cultivars that can perform well in marginal and water-stressed environments as their ability to reach water and or nutrients at greater depths is a key issue (see chapter: Breeding for Drought Tolerance). The wheat plant has two types of roots, the seminal (seed) roots and roots that initiate after germination, the nodal (crown or adventitious) roots. About six root primordia are present in the embryo. At germination, the primary root bursts through the coleorhiza, followed by the emergence of four or five lateral seminal roots. These form the seminal root system, which may grow to 2 m in depth and support the plant until the nodal roots appear. Nodal roots are associated with tiller development and are usually first seen when the fourth leaf emerges and tillering starts. Compared with the seminal roots, they are thicker and emerge more or less horizontally; when they first appear they are white and shiny (the white root stage). Nodal roots occur on the lower three to seven nodes (depending on environmental conditions and final number of leaves on the shoot). The uppermost node, on which roots occur, at the base of the culm, may be above soil level, and the roots may not penetrate the soil but appear as short pegs protruding from the stem. At maturity, the root system extends to between 1 and 2 m deep or more depending on soil conditions. Most roots occur in the top 30 cm of soil (Kirby E.J.M., 2002 from Botany of wheat plant FAOBW p22).

IV. Vernalization and Photoperiod (Taken from E. Acevedo et al. 2002 p.42
FAOBW)

Vernalization Wheats, which are responsive to vernalization, flower after the completion of a cold period. The double ridge stage is not reached until chilling requirements are met, and the vegetative phase is prolonged generating a higher number of leaves in the main shoot; the phyllochron, however, is not affected (Mossad et al., 1995). Two major flowering types of wheat are differentiated by their response to vernalization (Flood and Halloran, 1986): Spring-type wheat has a very mild response or no response at all to vernalization, and frost resistance is low. Winter-type wheats have a strong response to vernalization and require a period of cold weather to flower. In the early stages of growth, they are very resistant to frost (-20C), but frost resistance is gradually lost towards heading

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Fig. 8 Grain Development B

Integulment

Ovule
Antipodals

Central Cell with 2 polar nuclei

Synergids

Egg Cell

funiculus

A
micropyle

A) Diagram of pollen and ovule B) From top to bottom embryo development at 2, 7, 15, and 26 days after anthesis. ce = cellular endosperm em = embryo nu = nucellus sc = scutellum sp = shoot pole rp = root poll C) Grain at 6, 8, and 10 days after fertilization

Pollen Grain

Vegetative nucleus

Generative nucleus

Cytoplasm

Photos: 'WHEAT:THE BIG PICTURE'

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and flowering. The vernalization requirements of winter types may be fully substituted by short days at nonvernalizing temperatures between 21 and 16C (Evans, 1987). Flood and Halloran (1986) point out that vernalization may occur at three stages of the growing cycle of the wheat plant: during germination, during vegetative plant growth (GS1) and during seed formation in the mother plant. The effectiveness of low temperatures to accomplish vernalization decreases with increasing plant age, being almost nil after three months (Chujo, 1966; Leopold and Kriederman, 1975). Vernalization occurs at temperatures between 0 and 12C (Ahrens and Loomis, 1963; Trione and Metzger, 1970). Spring genotypes usually require temperatures between 7 and 18C for 5 to 15 days for floral induction, while winter types require temperatures between 0 and 7C for 30 to 60 days (Evans et al., 1975). Manupeerapan et al. (1992) observed that vernalization in winter genotypes stimulated cell division, overcoming an inhibitory process that occurs at high temperatures. Photoperiod After vernalization is completed, genotypes, which are sensitive to photoperiod, require a certain day-length to flower. Sensitivity to photoperiod differs among genotypes. Most cultivated wheats, however, are quantitative long-day plants. They flower faster as the day-length increases, but they do not require a particular length of day to induce flowering (Evans et al., 1975; Major and Kiniry, 1991). Stefany (1993) observed a period of insensitivity to day-length in wheat, which starts with germination. During this period, the plant develops foliar primordia only. This may be considered a juvenile phase, which is longer in winter wheat. The photoperiod is sensed by mature leaves and not by apical meristems (Barcell et al., 1992; Bernier et al., 1993). A single leaf is usually enough to sense the photoperiod for floral induction. Once the photoperiod insensitive period ends, floral induction starts and the reproductive stage begins (double ridge). The shorter the length of the day, the longer the inductive phase is (Major, 1980; Boyd, 1986), the longer the phyllochron (Cao and Moss, 1989a, 1989b; Mossad et al., 1995) and the bigger the flag leaf (Mossad et al., 1995). On the contrary, longer days advance floral induction (Evans et al., 1975). The development of the inflorescence after induction occurs at a rate that is also dependent on daylength in those genotypes sensitive to photoperiod (Stefany, 1993). The shorter the day, the longer the phase is from double ridge to terminal spikelet (Figure 3.2), increasing the period to terminal spikelet and the number of spikelets per spike. Changes in daylength after the terminal spikelet have no effect on floret initiation or anthesis date. Wheat adaptation to a wide range of latitudes occurs at lower levels of photoperiod sensitivity such that flowering is not retarded significantly if the day-length is shorter than optimal (Santibaez, 1994). Vernalization and photoperiod constitute the basic processes of the adaptation of wheat to various environments. Knowledge and genetic manipulation of them should continue to provide significant tools for adaptation and yield.

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V. Physiological Processes Driving Growth and Development

(Taken from E. Acevedo et al. 2002 p45 to 48)

The net carbon dioxide (CO2) assimilation at the tissue level constitutes the basis for growth. Many factors affect the net assimilation of CO2, among others, the growth and development stage of the plant and environmental characteristics, such as light, nitrogen, temperature, CO2 and water status. Four main basic processes are involved in photosynthesis: (i) a photochemical process determining the quantum yield and depending on light intensity; (ii) a biochemical process particularly linked to carboxylation; (iii) physico-chemical processes of CO2 transfer from the external air to the carboxylation sites; and (iv) the photorespiration process in C3 plants. At optimum temperature (20 to 25C), the maximum light saturation rates of photosynthesis (Amax) at the leaf level in bread wheat are between 15 and 25 mol CO/m2 s (25 to 40 mg CO2/dm2 h). Ninety percent of the light saturation rate is reached at 1 000 mol quanta/m2 s of photosynthetically active radiation (PAR). Wild relatives of wheat, however, may have substantially higher Amax than cultivated wheat (Austin, 1990). Much attention has been given to the question of how to increase total photosynthetic yield. Of the two photosynthetic parameters, quantum yield (rate of photosynthetic assimilation/incident light intensity) and Amax, a much greater improvement in canopy photosynthesis could be theoretically achieved by increasing the quantum yield. Unfortunately, the quantum yield of the photosynthetic process itself is very constant among genotypes (Austin, 1990). An improved discrimination of the enzyme ribulose 1,5 carboxylase oxygenase (rubisco) for CO2 with respect to oxygen (O2) would increase the quantum yield of the overall process by decreasing photorespiration (normally 25 percent of the energy produced by photosynthesis), but not much variation in the discrimination of rubisco has been found between species (Sommersville, 1986; Loomis and Amthor, 1996). Some scope appears to exist for selecting genotypes with a reduced maintenance respiration, which normally uses 2 to 3 percent of the dry weight per day (Robson, 1982), but its effect on radiation use efficiency would be low (Loomis and Amthor, 1996). Amax varies significantly among species and cultivars. In wheat, it has been known for some time that certain diploid ancestor species have higher Amax values than present advanced lines of bread and durum wheats (Dunstone et al., 1973); however, little progress has been made with respect to yield increases by this approach. Canopy photosynthesis Canopy photosynthesis is closely related to the photosynthetically active (400 to 700 mm) absorbed radiation (PARA) by green tissue in the canopy (Fischer, 1983). The PARA can be calculated from the fraction of solar radiation at the top of the canopy, which is transmitted to the ground (I/I0), such that: (2) PARA = RS * 0.5 * 0.9 * (1 - I/I0) where RS refers to the total solar radiation (MJ/m2 d); the factor 0.5 refers to the fraction of total solar energy, which is photosynthetically active; (1 - I/I0) is the fraction of total solar radiation flux, which is intercepted by the crop; and 0.9 * (1 - I/I0) is the fraction of

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radiation absorbed by the crop allowing for a 6 percent albedo and for inactive radiation absorption (Loomis and Amthor, 1996). The I/I0 essentially changes as the crop leaf area index (LAI) increases, and it is not very dependent on other factors, such as cloudiness or time of day. It is measured with a PAR sensor since the attenuation of RS in the canopy differs from that of PAR. The relationship between I/I0 and LAI fits a negative exponential (similar to the Beer Lambert Law), such that:

where e is the base of the natural logarithm and K is known as the canopy extinction coefficient. The canopy extinction coefficient in wheat crops ranges from 0.3 to 0.7 and is highly dependent on leaf angle (low K for erect leaves). From equation 3, it can be calculated that 95 percent PAR interception requires a LAI as high as 7.5 for erect leaves but a LAI of only about 4.0 for more horizontal leaves. The total canopy net photosynthesis is linearly related to PARA and so is crop growth rate (CGR, g/m2 d), which is the net accumulation of dry weight, such that: (4) CGR = RUE * PARA where RUE is the radiation use efficiency (g/m2 d). Final yield is therefore the product of cumulative seasonal radiation absorption, RUE and the portion of total biomass that goes to yield (harvest index). Potential radiation use efficiency in strong light depends on several factors: adequate water to allow high stomatal conductance and transport of CO2 into leaves; leaf arrangement relatively vertical to the radiation beam; good leaf nutrition to support large photosynthetic capacity; an active Benson-Calvin cycle to incorporate CO2; and appropriate canopy ventilation supplying CO2 and dissipation of heat (dissipation of excess energy due to light saturation). Due to environmental constraints, a quantum requirement of 10 mol quanta/mol CO2 under light-limited conditions may increase to 20 and 30 mol quanta/mol CO2 under field conditions with a decrease in RUE from 8.2 to 3.7 and 2.2 g/MJ PAR (Loomis and Amthor, 1996). Practical estimates of maximum RUE by these authors were 3.8 g/MJ PAR, which would occur with long cool days and moderate radiation (20 MJ/m2 d). Warm temperature, the small concentration of CO2 relative to O2 and light saturation limit attainment of a greater RUE. Measured values of RUE in a wheat crop are close to 3.0 g/MJ PARA when roots are included (Fischer, 1983). The RUE varies as Amax changes. Increases in the nitrogen of the canopy increase Amax and RUE. Frost at night and temperatures below 15C during the daytime can reduce Amax. Water stress has a small effect on RUE, but radiation intensity beyond a given value may reduce RUE. The RUE declines during grainfilling probably due to sink limitation and/or leaf senescence (Fischer, 1983). Most studies show no difference in CGR between genotypes, even when Amax varies (Austin et al., 1986), but a higher CGR at anthesis was related to higher yield in Australian modern wheat cultivars grown under water stress (Karimi and Siddique, 1991). A number of possibilities for utilizing the variability found for these physiological processes will be discussed further in the chapters: Breeding for Yield, and Breeding for Drought resistance. - 18 -

Questions to Test Understanding: 1. Explain the Zodoks decimal growth stages. 2. What are the disadvantages of tilling? 3. Define anthesis. 4. What takes place during the medium milk stage? 5. When does vernalization occur? 6. How does photoperiod affect inflorescence?

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References:
Ahrens and Loomis, 1963 Austin, 1990 Austin et al., 1986 Barcell et al., 1992 Boyd, 1986 Bernier et al., 1993 Cao and Moss, 1989a, 1989b Chujo, 1966
Duggan, B.L., R.A. Richards, A.F. van Herwaarden, and N.A. Fettell. 2005a. Agronomic evaluation of a tiller inhibition gene (tin) in wheat. I. Effect on yield, yield components, and grain protein. Australian Journal of Agricultural research 56: 169-178. Duggan, B.L., R.A. Richards, and A.F. van Herwaarden. 2005b. Agronomic evaluation of a tiller inhibition gene (tin) in wheat. II. Growth and partitioning of assimilate. Australian Journal of Agricultural Research 56: 179-186.

Dunstone et al., 1973

Taken from E. Acevedo et al. 2002 p.42 FAOBW

Evans, 1987
Evans L.T., J. Bingham, P. Johnson, and J. Sutherlands. 1972. Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears. Annals of Applied Biology 70:6776.

Evans et al., 1975 Fischer, 1983 Flood and Halloran (1986) Haun, 1973 Karimi and Siddique, 1991 Kirby E.J.M., 2002 from Botany of wheat plant FAOBW p22 Krumm et al., 1990 FAOBW44 Large, 1954 Leopold and Kriederman, 1975 Loomis and Amthor, 1996 Major, 1980 Manupeerapan et al. (1992 Miralles and Slafer 2007 Mossad et al., 1995 Percival, 1921 FAOBW 34 Robson, 1982 Santibaez, 1994 Siddique et al., 1989FAOBW44 Sommersville, 1986 Stefany, 1993 Trione and Metzger, 1970 Zadoks et al., 1974 FAOBW 44

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