See

discussions, stats, and author profiles for this publication at:

https://www.researchgate.net/publication/234176000

Dry matter accumulation and nutrient

uptake in flue-cured tobacco (Nicotiana
tabacum L.)

Article in Field Crops Research November 2004

DOI: 10.1016/j.fcr.2004.11.002

CITATIONS READS

22 398

2 authors, including:

Nicolaos K. Moustakas
Agricultural University of Athens
38 PUBLICATIONS 815 CITATIONS

SEE PROFILE

All content following this page was uploaded by Nicolaos K. Moustakas on 16 July 2017.

The user has requested enhancement of the downloaded file.

 Field Crops Research 94 (2005) 113
www.elsevier.com/locate/fcr

Dry matter accumulation and nutrient uptake in

flue-cured tobacco (Nicotiana tabacum L.)
N.K. Moustakasa,*, H. Ntzanisb
a
Agricultural University of Athens, Soil Science and Agr. Chemistry Lab., Iera Odos 75, Botanikos 118 55, Athens, Greece
b
Experimental Tobacco Station, Heraklitou 12, Agrinio 301 00, Greece
Received 2 November 2004; accepted 8 November 2004

Abstract

In a two-year field experiment using flue-cured tobacco, carried out in Agrinio (western Greece), dry matter accumulation
(DMA) was studied in addition to the uptake of the nutrients nitrogen (N), potassium (K), phosphorous (P), calcium (Ca), and
magnesium (Mg) on a weekly basis during the period from transplant to harvest over two cultivation seasons. Whole plants were
sampled and divided into leaves, stalks and roots. These were dried, weighed and DMA and the nutrient uptake determined. Both
DMA and nutrient uptake in plant parts as well as in whole plants follows a sigmoidal curve, accurately described by a logistic
equation. During growth there is a period when DMA and nutrient uptake in plant parts occurs at an intense rate. The time of
onset of this period and its duration varies with different plant parts. Maximum daily DMA occurs when 50% of the maximum
plant DMA has been achieved. Maximum daily nutrient uptake in aerial parts of the plant occurs approximately 1 week prior to
the maximum daily DMA. The period of rapid DMA and nutrient uptake in flue-cured tobacco coincides with the knee-high and
budding (rapid qrowth and elongation) stage (4175 DAT). Consequently during this period, the soil must have sufficient
nutrients available to supply the needs of the plant.
# 2004 Elsevier B.V. All rights reserved.

Keywords: Logistic equation; Rate of dry matter accumulation; Rate of nutrient uptake; Daily dry matter accumulation; Daily nutrient uptake

1. Introduction Corrective fertilisation between 35 and 40 DAT, a

The life cycle of flue-cured tobacco is relatively
short, ranging from 90 to 120 days and hence any
window of opportunity for intervention to correct any
nutrient deficits is brief (Miner and Tucker, 1990).
* Corresponding author. Tel.: +30 210 5294099;
fax: +30 210 5294092.
E-mail address: nmoustakas@aua.gr (N.K. Moustakas).
stage when the plants are adapting to a new
environment, is impractical due to the large size of
the plants.
In order for a rational fertilisation programme to be
developed, in addition to the nutrient status of the soil
in which the tobacco will be cultivated, the following
information is required: (a) the maximum nutrient
uptake determining the maximum yield, (b) the rate of
nutrient uptake throughout the life span of the plant to

0378-4290/$ see front matter # 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2004.11.002
2 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
determine the period of maximum nutrient uptake and
(c) the distribution of nutrients within the plant parts
during (leaves, stalks and roots) maturation in order to
determine the quantity of nutrients that will be
removed from the field upon harvest
Nutrient demands are estimated from knowledge of
the quantities of these nutrients removed by the plants
during a cultivation season in relation to the DMA,
largely in the aerial parts. The total DMA and the
quantity of nutrients taken up vary with the type of
tobacco, the fields residual nutrient status, planting
density, irrigation, climate and other environmental
factors (Collins and Hawks, 1993; Ceotto and Castelli,
2002). Consequently, nutrient uptakes reported in the
literature reflect the particular local cultural practices
and climatic conditions under which the tobacco is
grown (McCants and Woltz, 1967). Generally, Burley
and Maryland tobaccos have high demands for N and
are grown in fertile, medium to light textured soils.
Flue-cured tobacco is less demanding and will
satisfactorily grow in sandy and loamy-sand soils,
whilst orient tobaccos are grown on soils low in N
(Flower, 1999). Data for flue-cured tobacco, both
on DMA and nutrient uptake, under Mediterranean
conditions are scarce and concern only to the aerial
portion of plants with little or no reference to DMA
and nutrient uptake in the roots.
The purpose of this study was the investigation of
the patterns of DMA and the uptake of N, P, K, Ca and
Mg in leaves, stalks and roots of flue-cured tobacco
grown under Mediterranean conditions.
1.1. Theoretical considerations
The application of mathematics to the study of
plant growth rates has led to such a plethora of
research that it is now itself a specialised discipline of
plant physiology in its own right. The growth of a crop
passes through three consecutive phases which in their
entirety determine a quantitative relationship between
production/yield and time. This relationship is
described by the sigmoidal growth curve.
The logistic equation (Hunt, 1982)
a automatic weather station installed 40 m from the
W (1)
1 ebct
is a mathematical expression describing the sigmoidal
growth curve, i.e. the growth of a plant, where W is the
plant variable (namely DMA or nutrient uptake) at
time after transplanting t, where a is the maximum
value of DMA or nutrient uptake, b the initial DMA or
nutrient uptake, c an accumulation or uptake constant,
determined by calculation. The logistic equation
expresses DMA as a factor of time. Thence the growth
rate, i.e. increase in production per unit time, is
calculated from the first derivative of Eq. (1) as:
dW ab ebct
(2)
dt 1 ebct 2
Half the maximum value of the result of Eq. (1) gives
the point of inflexion and corresponds to the time t at
which the maximum growth rate is observed.
Of interest is the second derivative of Eq. (1)
dW 2 ac ebct ebct  1
(3)
d2 t 1 ebct 3
The value of t at which the second derivative equals
zero also corresponds to the time of maximum growth
rate.
2. Materials and methods
For the purposes of this investigation, a field
experiment was carried out over 2 years in Agrinio,
western Greece (3883700 N, 2182300 E, 45 m above sea
level). The soil of the site is a clayey loam (CL) (fine,
mixed, thermic, typic xerofluvent) with 26% sand,
43% silt and 31% clay, pH 7.1, cation-exchange
capacity (CEC) of 20.4 cmolc kg1 soil, organic
matter 1.8%, exchangeable Ca, Mg and K of 17.2,
1.7 and 0.5 cmolc kg1, respectively, and available P
(P Olsen) 6.2 mg kg1 soil. Prior to this study the
field was under corn cultivation (1993).
The climate of the region is classed as a thermic
Mediterranean (FAO-UNESCO, 1973). Average rain-
fall (for 30 years) is 1022 mm with 68% falling
between November and April, with an average air
temperature of 14.7 8C. Climatic data such as air
temperature, air humidity, global radiation, wind
velocity and precipitation were measured from an
experimental site. During the cultivation periods of
1994 and 1995, maximum air temperature was greater
than 30 8C in JuneAugust, whilst the minimum
temperature ranged from 13 to 19 8C and 11 to 18 8C
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
for 1994 and 1995, respectively, over the same period.
No difference in rainfall distribution during the two
experimental periods was observed. Briefly rainfall
was 98 mm in 1994 distributed as 23% in May, 39% in
July and 38% in August. In 1995, rainfall was 101 mm
distributed as 24% in May, 35% in July and 41% in
August. Tobacco plants were irrigated 12 times in
1994 and 10 times in 1995 with a total irrigation of
350 mm each cultivation period. The quantity of water
used for irrigation was equal to the maximum
evapotranspiration rate (ETc) which was estimated
from the potential evapotranspiration (ETp) and the
crop constant for tobacco (Kc). ETp was calculated
using the minimum and maximum air temperatures,
duration of solar radiation, relative humidity and wind
speed, using the Penman Equation (1948) as modified
by Frere (1979). The value of the constant Kc for
tobacco during the growing season was taken as 0.4
for the recovery period, 0.70.8 for the phase of rapid
growth and elongation, 0.91.0 for the stage of
flowering and topping and finally 0.750.85 for the
maturation and seed formation phase (Doorenbos and
Pruit, 1977). Irrigation was carried out when the
moisture content at 0.3 m was less than 40% of the
available water moisture. Soil moisture was measured
using tensiometers installed in the field.
The experimental field comprised 18 plots
(150 m2) arranged in 3 rows of 6 plots with a distance
of 1.5 m between rows and 1 m between plots. On 18th
May 1994 and 19th May 1995, each plot was planted
up with flue-cured tobacco plants (McN-944) in 15
rows, each row 10 m long, with a distance of 1 m
between rows and 0.5 m between plants (20,000 plants
per ha). Half of the plot was used to monitor DMA and
nutrient uptake, the other half being used for
collection and processing of mature tobacco leaves
to determine crop yield. Fertilisation was carried out
each year using quantities appropriate for maximum
yield and quality production of flue-cured tobacco.
The total amounts of fertiliser used were 6 g N m2,
12 g P m2, and 24 g K m2. Applications were made
23 days before transplanting using 4 g N m2,
12 g P m2 and 17 g K m2 using calcium ammonium
nitrate (26% N), ordinary superphosphate (21% P2O5)
and potassium sulphate (50% K2O). One month after
transplanting, an application of 2 g N m2 and
7 g K m2 was applied in the form of KNO3 (13%
N and 46% K2O). The fertiliser was uniformly
3
broadcast over the soil surface of each plot before
being incorporated into the soil to a depth of 0.15
0.2 m by hand. Topping took place when 50% of the
plants in each plot were at full bloom (75 and 76 DAT
in 1994 and 1995, respectively) and to a height such
that 2224 leaves remained. All cultivation practices
took place in strict accordance with the guidelines of
the National Tobacco Institute of Greece (1996).
On an approximately weekly basis, three plants
were randomly selected from each experimental plot
from 34 DAT until 96 DAT (full maturation and seed
production) resulting in the sampling of 54 plants per
sampling date. The length of the various stages of
tobacco development and the number of samplings
during that period (National Tobacco Institute of
Greece, 1996) are shown below:
 recovery (adaptation): 3035 days, 1 sampling;
 knee-high: 1015 days, 2 samplings;
 budding (rapid growth and elongation): 1015 days,
2 samplings;
 flowering and topping, beginning of harvest: 2535
days, 2 samplings;
 maturation and seed formation: 2025 days, 2
samplings.
Each plant was extracted from the soil by digging a
trench around it 0.3 m2 by 0.4 m deep and removing it
as a block. The roots were washed well to remove all
traces of soil and the plants then separated into leaves,
stalks and roots. Individual parts were then washed to
remove all traces of soil. The corresponding parts of
the three plants sampled were combined to produce a
bulk sample for each plant part and each experimental
plot. For statistical purposes, there were thus 18 re-
plicates. The fresh weight of the bulk sample plant
parts was recorded and the samples were then dried to
a constant weight in an oven at 70 8C whereon dry
weight was then recorded. The parts were then ground
to pass a 250 mesh sieve. A portion of the ground
samples was incinerated at 550 8C and the ash diss-
olved in concentrated nitric acid. This solution was
used for the determination of Ca, Mg, P and K. Co-
ncentrations of Ca and Mg were determined using
atomic absorption spectrometry with an acetylene
N2O and acetyleneoxygen flame for Ca and Mg,
respectively. Potassium was determined using flame
photometry. The ascorbic acid method of Murphy
4 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
Rilley was used to determine P. Total nitrogen was
determined using the Kjeldahl method. The analysis of
plant parts took place in accordance with the methods
reported by Page (1982).
The dry weight of leaves, stalks and roots was
determined for each sampling date and expressed as
g m2. The amount of N, K, P, Ca, and Mg taken up
and distributed within plant parts was calculated by
multiplying the dry weight by the corresponding
elemental concentration and expressed as g m2. The
leaves of plants not sampled for elemental analysis
were harvested and dried in a bulk-curing furnace,
appropriately treated and the tobacco yield calculated
per hectare.
After data collection, statistical analyses were
performed in the following order to:
(a) examine the effect of year and date on DMA and
nutrient uptake and distribution in whole plant and
plant parts;
(b) determine whether data for the 2 years could be
combined and analysed, using Bartletts x2 test for
homogeneity;
(c) examine the adaptation of our data using non-
linear regression to a logistic equation and
estimation of the parameters a, b, and c;
(d) statistically evaluate predicted and measured
values.
3. Results and discussion
The distribution of rainfall, as well as other factors
influencing tobacco growth, such as daily minimum
and maximum air temperatures, was satisfactory
in both years. The average annual production of
processed leaves was 3450 kg ha1 (380.3).
An examination of the data showed that experi-
mental year and sampling date had no effect on DMA
nor nutrient uptake and distribution. Bartletts x2 test
showed that combining the data from both years was
acceptable. In addition, deviation in the transplant date
in the two experimental years was almost non-existent
and hence for any given sampling date the plants were
at the same stage of development in each year. In the
analysis that follows, all values given are the averages
of the data for the 2 years combined.The DMA in
leaves (DML), stalks (DMS), roots (DMR), aerial
parts (DMAe = DML + DMS) and whole plant (DMP)
during the growth period follows a sigmoidal curve
(Figs. 15) which can be accurately described by the
logistic equation:
a
W
1 ebct
where W is the accumulation of DML, DMS, DMR,
DMAe and DMP at a time t, a is the maximum value of
DMA, b the initial DMA, c an accumulation constant,
t the time in days after transplanting. The values for a,
b, and c were calculated using Rosenbrocks method
(Machura and Mulawa, 1973) with the statistical soft-
ware application Statistica for Windows (StatSoft,
1995). The correlation coefficient r between predicted
and observed values is very high (Table 1) and ensures
the statistical integrity of the curve.
Figs. 15 show the accumulation curves for whole
plants and plant parts resulting from the application of
the logistic equation to the data. A study of these
curves gives us the following information.
3.1. Dry matter accumulation
Dry matter accumulation in whole plants and plant
parts was relatively slow until 35 DAT, indicating the
slow adaptation of the plants to their new environment.
Similar periods of adaptation have been reported by
Raper and McCants (1967), Atkinson et al. (1977),
and Mylonas and Pangos (1980) for flue-cured, Burley
and orient type tobaccos, respectively.
At maturation in flue-cured plants 848 g DM m2
accumulated distributed as 356 g m2 in leaves,
277.2 g m2 in stalks and 240.49 g m2 in roots
(Fig. 5) representing 42.3%, 33% and 28.5%,
respectively, of total DM. The sum of DM in plant
parts is not equal to the total DMA as a result of
smoothing applied during the regression analysis.
In the period between 34 and 96 DAT, between
transplant and maturation, there is rapid DMA in plant
parts and whole plants (Figs. 1a, 2a, 3a, 4a and 5a).
This period differs between plant parts both in respect
to onset as well as to duration. In particular, in leaves
this period of rapid DMA is between 41 and 75 DAT
(Fig. 1a) in which 93% of total DMA is achieved,
corresponding to 39.4% DMP. Similarly, Suzuki et al.
(1970) report rapid changes in DMA in Burley tobacco
leaves from 30 to 75 DAT. In stalks, there is a period of
rapid DMA from 48 to 90 DAT (Fig. 2a) where 95% of
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113 5

Fig. 1. (a) Dry matter accumulation and nutrient uptake curves in leaves of flue-cured tobacco. (b) Leaf growth rate and nutrient uptake rate
curves of flue-cured tobacco.

DMS is achieved, corresponding to 11.4% of DMP.
This period of rapid DMA occurs in roots from 48 to
90 DAT (Fig. 3a) where 82% of DMR is achieved,
corresponding to 11.8% of DMP.
From transplant to 48 DAT (i.e. the first half of
the period from transplant to maturation) DMA is
86.6 g m2 in leaves, 18 g m2 in stalks and 3 g m2
in roots, representing 23%, 6.5% and 1.3% of DMA in
leaves, stalks and roots or 9.0%, 2.0% and 0.4%,
respectively, of DMP at maturation.
DMA in leaves, stalks and roots per unit time is
calculated from Eq. (2) and expresses the rate of
growth, i.e. the daily DMA in the various parts of the
tobacco plant. The maximum daily DMA differs
between plant parts both in value and in date of onset.
The maximum DMA in leaves, stalks and roots is
observed on 56, 70 and 69 DAT, at 89.5, 72.4 and
60.2 g m2, respectively (Figs. 1b, 2b and 3b).
The rate of increase in DMP rises until 62 DAT
when it reaches its maximum, after which it declines
to zero when DMA is at its maximum. Fig. 6 shows
that the maximum growth rate, i.e. maximum daily
DMA, is achieved at time t where the second factor of
Eq. (3) tends to zero and DMA is at half its maximum.
The same pattern in daily DMA is observed in plant
parts for flue-cured tobacco. It is noteworthy that the
6 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113

Fig. 2. (a) Dry matter accumulation and nutrient uptake curves in stalks of flue-cured tobacco. (b) Stalk growth rate and nutrient uptake rate
curves of flue-cured tobacco.

maximum daily DMA in leaves, stalks and roots
appears towards the end of the budding stage (early
floweringtopping stage).
3.2. Nutrient uptake
The uptake of nutrients in whole flue-cured tobacco
plants, as well as the uptake and distribution in
individual plant parts follows a sigmoidal curve
(Figs. 15) described by the logistic equation:
a and 68 DAT where 85% of whole plant N is taken up.
U
1 ebct
where U is nutrient uptake (g m2) at time t, a is
maximum nutrient uptake (g m2), b the initial nutri-
ent uptake (g m2), c a nutrient uptake constant, t time
in days after transplant and e the natural logarithm.
Again the sum of uptakes is not equal to the total
uptake due to smoothing in the regression analysis.
3.2.1. Nitrogen uptake
The pattern of N uptake in whole plants and in plant
parts is shown in Figs. 1a, 2a, 3a, 4a and 5a. A period
of rapid change in N uptake is observed between 41
The maximum daily uptake of N is observed on 55
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113 7

Fig. 3. (a) Dry matter accumulation and nutrient uptake curves in roots of flue-cured tobacco. (b) Root growth rate and nutrient uptake rate
curves of flue-cured tobacco.

DAT at 4.5 g m2 (Fig. 5b). In leaves, the period of
rapid change in uptake occurred between 41 and 68
DAT (Fig. 1a) where 93% of total leaf N was taken up,
corresponding to 56% of whole plant N. The
maximum daily N uptake distributed in leaves was
found on 54 DAT at 2.7 g m2, in stalks 67 DAT at
0.86 g m2 and in roots 64 DAT at 1.1 g m2 (Figs. 1b,
2b and 3b). The amount of N in the whole plant at
maturity was 18.24 g m2, distributed as 10.9 g m2
in leaves, 3.22 g m2 in stalks and 4.6 g m2 in roots,
representing 60%, 18% and 25% whole plant N,
respectively (Fig. 5a). During the growth period from
transplant to 48 DAT (i.e. the first half of the growth-
maturation phase) N taken up and distributed in
leaves, stalks and roots was 2.9, 0.5 and 0.13 g m2
corresponding to 16%, 2.6% and 0.7% whole plant N,
respectively. During the floweringtopping stage, the
leaves, stalks and roots contained 10.7, 2.3 and
4.22 g N m2, respectively. These values represent
98%, 72% and 93% of total uptake in leaves, stalks
and roots, respectively, and correspond to 58%, 13%
and 23% of whole plant N uptake. According to
8 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113

Fig. 4. (a) Dry matter accumulation and nutrient uptake curves in the aerial parts of flue-cured tobacco. (b) Crop growth rate and nutrient uptake
rate curves of aerial parts of flue-cured tobacco.

McCants and Woltz (1967), the uptake and distribu-
tion of small quantities of N to the leaves of flue-cured
tobacco after flowering is desirable to ensure good
quality.
3.2.2. Potassium uptake
A period of rapid change in K uptake is observed
between 41 and 68 DAT where 81% of whole plant K
is taken up (Fig. 5a). The maximum daily uptake of K
is observed on 61 DAT at 4.6 g m2 (Fig. 5b). In
leaves, the period of rapid change in uptake occurred
between 41 and 68 DAT, in stalks between 41 and 90
DAT and in roots between 48 and 75 DAT (Figs. 1a, 2a
and 3a) where 93%, 92% and 93% of total leaf, stalk
and root K was taken up, corresponding to 18%, 33%
and 22.5% of whole plant K. The amount of K in the
whole plant at maturity was 18.54 g m2, distributed
as 7.6 g m2 in leaves, 6.7 g m2 in stalks and
5.3 g m2 in roots, representing 41%, 36% and 29%
whole plant K, respectively. These values represent
98%, 62% and 93% of total K uptake in leaves, stalks
and roots, respectively, and correspond to 40.5%, 23%
and 27% of whole plant K uptake. During the growth
period from transplant to 48 DAT (i.e. the first half of
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113 9

Fig. 5. (a) Dry matter accumulation and nutrient uptake curves over a growing season of flue-cured tobacco in whole plants. (b) Crop growth rate
and nutrient uptake rate curves over a growing season of flue-cured tobacco.

the growth-maturation phase) K taken up and
distributed in leaves, stalks and roots was 1.9, 0.7
and 0.14 g m2 corresponding to 10.3%, 3.9% and
0.8% whole plant K, respectively.
The distribution of K in tobacco plants indicates
that K is essential even after topping. McCants and
Woltz (1967) report that application of a K containing
fertiliser 34 weeks after transplanting favours the
production of good tobacco. The maximum daily K
uptake distributed to leaves was found on 54 DAT at
1.91 g m2 (Fig. 1b) showing that K must be present in
significant quantities even at the early stages of
development. The maximum daily K uptake distrib-
uted in stalks occurred 73 DAT at 1.86 g m2 and in
roots 64 DAT at 1.33 g m2 (Fig. 1b)
The quantities of N and K taken up and distributed
in aerial parts of the plant are approximately the same
at 18.5 g m2. McCants and Woltz (1967) report that
while flue-cured tobacco takes up greater quantities of
K than N, greater quantities of K-fertiliser must be
added for the further reason that a high K uptake
improves tobacco quality.
3.2.3. Calcium uptake
A period of rapid change in Ca uptake is observed
between 41 and 75 DAT where 90% of whole plant Ca
10 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113

Table 1 and root Ca was taken up, corresponding to 58%, 14%

Computation parameters a, b, c from the logistic equation using non-
and 4% of whole plant Ca. The amount of Ca in the
linear regression technique and correlation coefficient r between
observed and predicted values whole plant at maturity was 17.7 g m2, distributed as
9.6 g m2 in leaves, 2.3 g m2 in stalks and 3.9 g m2
Variable a b c r
in roots, representing 48.25%, 25% and 27% whole
DMLa 357.8 7.90 0.14 0.992
plant Ca, respectively. During the growth period from
DMS 289.73 8.59 0.12 0.996
DMR 240.68 15.85 0.24 0.97 transplant to 48 DAT (i.e. the first half of the growth-
DMAe 639.57 7.34 0.12 0.992 maturation phase) Ca taken up and distributed in
DMP 855.57 7.93 0.13 0.99 leaves, stalks and roots was 2.5, 0.3 and 0.17 g m2
NLb 10.88 9.69 0.18 0.973 corresponding to 16%, 2% and 1% whole plant Ca,
NS 3.43 6.97 0.09 0.997
respectively. During the floweringtopping stage, the
NR 4.56 14.21 0.22 0.97
NAe 13.7 9.01 0.16 0.992 leaves, stalks and roots contained 9.1, 1.9 and
N 18.27 9.06 0.16 0.996 3.2 g Ca m2, respectively. These values represent
KLc 7.68 10.29 0.19 0.887 95%, 83% and 81% of total uptake in leaves, stalks
KS 7.47 6.64 0.09 0.989 and roots, respectively, and correspond to 58%, 12%
KR 5.33 14.6 0.23 0.97
and 20% of whole plant Ca uptake. The maximum
KAe 13.27 8.93 0.15 0.992
K 18.57 9.73 0.16 0.996 daily Ca uptake distributed in leaves was found on 55
PLd 0.49 8.98 0.17 0.867 DAT at 2.4 g m2, in stalks 62 DAT at 0.6 g m2 and in
PS 0.25 13.2 0.22 0.939 roots 66 DAT at 0.98 g m2 (Figs. 1b, 2b and 3b).
PR 0.27 13.03 0.20 0.955 Calcium is third in order of importance of the
PAe 0.75 9.56 0.17 0.992
nutrients taken up in large quantities by Flue-cured
P 1.01 9.99 0.17 0.996
CaLe 9.62 8.06 0.15 0.996 tobacco. McCants and Woltz (1967) refer to the fact
CaS 2.36 7.98 0.13 0.999 that the major macronutrient for Flue-cured tobacco is
CaR 3.91 11.23 0.17 0.996 K, however the uptake and distribution of K and Ca to
Ca Ae 11.92 7.94 1.14 0.992 leaves is dependent on the nutrient status of the plant
Ca 15.84 7.74 0.13 0.996
and the availability of Ca (Elliot, 1978).
MgLf 1.54 7.99 0.14 0.992
MgS 0.66 7.59 0.11 0.995
MgR 0.87 12.9 0.20 0.986 3.2.4. Magnesium uptake
Mg Ae 2.17 7.49 0.13 0.992 A period of rapid change in Mg uptake is observed
Mg 3.06 8.12 0.13 0.996 between 41 and 75 DAT where 89% of whole plant Mg
a
DML, DMS, DMR, DMAe, DMP: dry matter accumulation in is taken up (Fig. 5a). The maximum daily uptake of
leaves, stalks, roots, aerial plant parts and in whole plants, respec- Mg is observed on 61 DAT at 0.8 g m2 (Fig. 5b). In
tively.
b leaves, the period of rapid change in uptake occurred
NL, NS, NR, NAe, N: nitrogen uptake in leaves, stalks, roots,
aerial plant parts and whole plants, respectively. between 41 and 75 DAT, in stalks between 41 and 90
c
KL, KS, KR, KAe, K: potassium uptake in leaves, stalks, roots, DAT and in roots between 48 and 90 DAT (Figs. 1a, 2a
aerial plant parts and whole plants, respectively. and 3a) where 95%, 96% and 99% of total leaf, stalk
d
PL, PS, PR, PAe, P: phosphorus uptake in leaves, stalks, roots, and root Mg was taken up, corresponding to 48%, 21%
aerial plant parts and whole plants, respectively.
e and 28% of whole plant Mg.
CaL, CaS, CaR, CaAe, Ca: calcium uptake in leaves, stalks,
roots, aerial plant parts and whole plants, respectively. The amount of Mg in the whole plant at maturity
f
MgL, MgS, MgR, MgAe, Mg: magnesium uptake in leaves, was 3.03 g m2, distributed as 1.54 g m2 in leaves,
stalks, roots, aerial plant parts and whole plants, respectively. 0.6 g m2 in stalks and 0.6 g m2 in roots, represent-
ing 50%, 21% and 28% of whole plant Mg,
is taken up (Fig. 5a). The maximum daily uptake of Ca respectively. During the growth period from transplant
is observed on 61 DAT at 3.9 g m2 (Fig. 5b). In to 48 DAT (i.e. the first half of the growth-maturation
leaves, the period of rapid change in uptake occurred phase) Mg taken up and distributed in leaves, stalks
between 34 and 75 DAT, in stalks between 41 and 90 and roots was 0.4, 0.1 and 0.002 g m2 corresponding
DAT and in roots between 48 and 90 DAT (Figs. 1a, 2a to 13%, 2% and 1% of whole plant Mg, respectively.
and 3a) where 95%, 98% and 98% of total leaf, stalk During the floweringtopping stage, the leaves, stalks
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
Fig. 6. Dry matter accumulation curve, crop growth rate curve (first derivative) and crop growth acceleration curve (second derivative) over a
growing season in whole plants of flue-cured tobacco.
and roots contained 1.4, 0.5 and 0.8 g Mg m2,
respectively. These values represent 95%, 70% and
90% of total uptake in leaves, stalks and roots,
respectively, and correspond to 48.16%, 16% and 26%
of whole plant Mg uptake. The maximum daily Mg
uptake distributed in leaves was found on 56 DAT at
0.38 g m2, in stalks 67 DAT at 0.17 g m2 and in
roots 64 DAT at 0.22 g m2 (Figs. 1b, 2b and 3b).
3.2.5. Phosphorous uptake
A period of rapid change in P uptake is observed
between 41 and 75 DAT where 96% of whole plant P is
taken up (Fig. 5a). The maximum daily uptake of P is
observed on 55 DAT at 0.24 g m2 (Fig. 5b). In leaves,
the period of rapid change in uptake occurred between
34 and 61 DAT, in stalks between 41 and 75 DAT and
in roots between 48 and 90 DAT (Figs. 1a, 2a and 3a)
where 94%, 95% and 99% of total leaf, stalk and root P
was taken up, corresponding to 46%, 24% and 26%
of whole plant Mg.
The amount of P in the whole plant at maturity
was 1.0 g m2, distributed as 0.5 g m2 in leaves,
0.25 g m2 in stalks and 0.27 g m2 in roots,
representing 49%, 25% and 27% of whole plant P,
respectively. During the growth period from transplant
to 48 DAT (i.e. the first half of the growth-maturation
phase) P taken up and distributed in leaves, stalks and
roots was 0.16, 0.01 and 0.01 g m2 corresponding to
16%, 25% and 26% of whole plant P, respectively.
11
During the floweringtopping stage, the leaves, stalks
and roots contained 0.5, 0.24 and 0.24 g P m2,
respectively. These values represent 98%, 96% and
90% of total uptake in leaves, stalks and roots,
respectively, and correspond to 49%, 25% and 26% of
whole plant P uptake. It is noticeable that P uptake
occurs during the first stages of development prior to
topping. The maximum daily P uptake distributed in
leaves was found on 52 DAT at 0.12 g m2, in stalks
61 DAT at 0.06 g m2 and in roots 64 DAT at
0.07 g m2 (Figs. 1b, 2b and 3b).
3.2.6. Aerial plant parts
The DMA in aerial parts of the plant exhibits a
period of rapid change between 48 and 90 DAT
(Fig. 5a). Up to 48 DAT 12% DMA was produced and
18% N, 13.4% K, 17.6% Ca, 15.3% Mg and 17.2% P
of the whole plant content has been taken up and
distributed in aerial parts. Atkinson et al. (1977),
working with Burley tobacco, report that only 18%
DMA of the aerial plant parts is produced during the
first half of the growth period. Grizzard et al. (1942),
working on flue-cured tobacco, report that during the
first half of a 90-day cultivation period, 18% of DMA
was apportioned to the aerial parts of the plant.
The maximum daily DMA in aerial parts of the
plants was observed 62 DAT at 159.9 g m2 (Fig. 4b).
The maximum daily uptake and distribution of N in
aerial parts was recorded 55 DAT at 3.43 g m2
12 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
(Fig. 4b). Maximum daily uptake of K, Ca, Mg and P
in aerial parts was observed 57, 56, 58 and 55 DAT at
3.13, 2.98, 0.54 and 0.19 g m2, respectively (Fig. 4b).
It is notable that the maximum daily uptake of all
nutrients in aerial parts took place 57 days prior to the
attainment of maximum DMA. Raper and McCants
(1967) report that in flue-cured tobacco, 50% of N and
K of the plant is distributed to aerial parts during the
early growth stage, specifically from 35 to 42 DAT and
that the uptake and distribution of Ca, Mg and P occurs
at a relatively steady rate throughout the growth
period. Atkinson et al. (1977), Bruns and McIntosh
(1988) and Bertinuson et al. (1970) working with
Burley, Maryland and shade-grown wrapper tobacco
respectively report that the daily uptake and distribu-
tion of nutrients to aerial parts was greater during the
later stages of development.
The distribution of nutrients was different in
tobacco plant parts as shown in Figs. 13. The greater
part of N (60%) and of Ca (59%) was distributed in
leaves, resulting in their removal from the field at
harvest. In contrast, the major portion of K taken up by
plants was distributed in stalks (36%) and roots (29%)
which theoretically are returned to the soil at the end
of the cultivation season. The distribution of Mg and P
was equal between those parts harvested (4849% in
leaves) and those remaining in the field after harvest
(roots and stalks).
4. Conclusions
During growth there is a period when DMA in plant
parts occurs at an intense rate. The time of onset of this
period and its duration vary with different plant parts.
Maximum daily DMA occurs when 50% of the
maximum plant DMA has been achieved.
During growth there is a period of rapid change in
nutrient uptake whose onset and duration vary not only
with regard to the time of onset and duration but also to
specific nutrient, and plant part. Maximum daily
nutrient uptake in aerial parts of the plant occurs
approximately 1 week prior to the maximum daily
DMA.
The period of rapid DMA and nutrient uptake in
flue-cured tobacco coincides with the knee-high and
budding (rapid qrowth and elongation) stage (between
41 and 75 DAT). Consequently during this period, the
soil must have sufficient nutrients available to supply
the needs of the plant.
The greater part of N and Ca taken up by flue-cured
tobacco is distributed in leaves (60%) meaning that a
substantial proportion of these nutrients is removed at
harvest and should be replaced by appropriate
fertilisation. In contrast, K, though taken up in equal
quantities as N is distributed throughout the plant
differently with 65% remaining in stalks and roots
after harvest.
Hence for coarse-textured soils a split application
of 1/3 N and K fertiliser 23 days prior to transplanting
and 2/3 2530 DAT is recommended. For fine-textured
soils 2/3 prior to transplanting with an additional
application of the remaining 1/3 2530 DAT is
recommended in order to fulfil the plants require-
ments.
Acknowledgement
The authors would like to thank Ms. Susan Coward
for her help in the preparation of this paper.
References
Atkinson, W.O., Bush, L.P., Sims, J.L., 1977. Dry matter and
nutrient accumulation in Burley tobacco. Tob. Sci. 21, 8182.
Bertinuson, T.A., Larssen, E., Teveris, B., Comfort, M., Petersen,
M., 1970. Nutrient uptake and dry matter accumulation of
Connecticut shade-grown wrapper tobacco for three consecutive
years. Tob. Sci. 14, 155157.
Bruns, H.A., McIntosh, M.S., 1988. Growth rates and nutrients
concentrations in Maryland tobacco. Tob. Sci. 32, 8287.
Ceotto, E., Castelli, F., 2002. Radiation-use efficiency in flue-cured
tobacco (Nicotiana tabaccum L.): response to nitrogen supply,
climatic variability and sink limitation. Field Crops Res. 74,
117130.
Collins, W.K., Hawks, S.N., 1993. Principles of Flue-cured Produc-
tion. N. C. State University, Raleigh, NC.
Doorenbos, J., Pruit, W.O., 1977. Crop water requirements. Irriga-
tion and Drainage. Paper 24 (revised). FAO, Rome.
Elliot, J.M., 1978. A survey of flue-cured tobacco grown in Ontario
in 1976. Part II. Nutrient elements. Lighter 47, 1921.
FAO-UNESCO, 1973. Carte bioclimatique de la zone Mediterranee.
O.N.M., Paris.
Flower, F.C., 1999. Field practices. In: Layten Davis, D., Nielsen,
M.T. (Eds.), Tobacco, Production, Chemistry and Technology.
Blackwell Science, Oxford, UK.
Frere, M., 1979. A Method for the Practical Application of the
Penman Formula for the Estimation of Potential Evapotranspira-
 N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113
tion and Evaporation from Free Water Surface. FAO, Rome,
AGP:Eco/1979/1.
Hunt, R., 1982. Plant Growth Curves. Edward Arnold, London.
Grizzard, A.L., Davies, H.R., Kangas, L.R., 1942. The time and
rate of nutrients absorption by flue-cured tobacco. Agron. J. 34,
327339.
Machura, M., Mulawa, A., 1973. Rosenbrock function minimization
Algorithm 450. Commun. ACM 16, 482483.
McCants, C.B., Woltz, W.G., 1967. Growth and mineral nutrition of
tobacco. Adv. Agron. 19, 211265.
Miner, G.S., Tucker, M.R., 1990. Plant analysis as an aid in
fertilizing tobacco. In: Westerman, R.L. (Ed.), Soil Testing
and Plant Analysis SSSA Book, Ser. 3. SSSA, Madison, WI,
pp. 645657.
Mylonas, V.A., Pangos, E.A., 1980. Dry matter and nutrient
accumulation in various plant parts of oriental neutral type
View publication stats
13
tobacco.. In: Proceedings of CORESTA Congress, 2024
November, Manila, The Phillipines.
National Tobacco Institute of Greece, 1996. Guidelines for Tobacco
Production (in Greek).
Page, A.L., 1982. Chemical and microbiological properties. In:
Page, A.L. (Ed.), Methods of Soil Analysis. Part 2. Agronomy
9.. ASA, Inc., Madison, WI.
Penman, H.L., 1948. Natural evaporation from open water, bare
soils s and grass. Proc. Roy. Soc. A 193, 120146.
Raper, C.D., McCants, C.B., 1967. Nutrient accumulation in Flue-
cured tobacco. Tob. Sci. 11, 190.
Statsoft Inc., 1995. Statistica for Windows. tatSoft, Inc., Tulsa, OK.
Suzuki, M., Shinochara, T., Kona, K., 1970. Studies on the growth
and the absorption of nutrients by Burley tobacco plants growth
on Ando soils (Kuruboku). Morioka Lab. Tob. Exp. Sta. Bull.
3, 91109.