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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

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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


stage when the plants are adapting to a new
The life cycle of flue-cured tobacco is relatively environment, is impractical due to the large size of
short, ranging from 90 to 120 days and hence any the plants.
window of opportunity for intervention to correct any In order for a rational fertilisation programme to be
nutrient deficits is brief (Miner and Tucker, 1990). 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
* Corresponding author. Tel.: +30 210 5294099;
fax: +30 210 5294092. uptake determining the maximum yield, (b) the rate of
E-mail address: nmoustakas@aua.gr (N.K. Moustakas). 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 plant variable (namely DMA or nutrient uptake) at
(c) the distribution of nutrients within the plant parts time after transplanting t, where a is the maximum
during (leaves, stalks and roots) maturation in order to value of DMA or nutrient uptake, b the initial DMA or
determine the quantity of nutrients that will be nutrient uptake, c an accumulation or uptake constant,
removed from the field upon harvest determined by calculation. The logistic equation
Nutrient demands are estimated from knowledge of expresses DMA as a factor of time. Thence the growth
the quantities of these nutrients removed by the plants rate, i.e. increase in production per unit time, is
during a cultivation season in relation to the DMA, calculated from the first derivative of Eq. (1) as:
largely in the aerial parts. The total DMA and the
dW ab ebct
quantity of nutrients taken up vary with the type of (2)
tobacco, the fields residual nutrient status, planting dt 1 ebct 2
density, irrigation, climate and other environmental Half the maximum value of the result of Eq. (1) gives
factors (Collins and Hawks, 1993; Ceotto and Castelli, the point of inflexion and corresponds to the time t at
2002). Consequently, nutrient uptakes reported in the which the maximum growth rate is observed.
literature reflect the particular local cultural practices Of interest is the second derivative of Eq. (1)
and climatic conditions under which the tobacco is
dW 2 ac ebct ebct  1
grown (McCants and Woltz, 1967). Generally, Burley (3)
and Maryland tobaccos have high demands for N and d2 t 1 ebct 3
are grown in fertile, medium to light textured soils. The value of t at which the second derivative equals
Flue-cured tobacco is less demanding and will zero also corresponds to the time of maximum growth
satisfactorily grow in sandy and loamy-sand soils, rate.
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 2. Materials and methods
conditions are scarce and concern only to the aerial
portion of plants with little or no reference to DMA For the purposes of this investigation, a field
and nutrient uptake in the roots. experiment was carried out over 2 years in Agrinio,
The purpose of this study was the investigation of western Greece (3883700 N, 2182300 E, 45 m above sea
the patterns of DMA and the uptake of N, P, K, Ca and level). The soil of the site is a clayey loam (CL) (fine,
Mg in leaves, stalks and roots of flue-cured tobacco mixed, thermic, typic xerofluvent) with 26% sand,
grown under Mediterranean conditions. 43% silt and 31% clay, pH 7.1, cation-exchange
capacity (CEC) of 20.4 cmolc kg1 soil, organic
1.1. Theoretical considerations matter 1.8%, exchangeable Ca, Mg and K of 17.2,
1.7 and 0.5 cmolc kg1, respectively, and available P
The application of mathematics to the study of (P Olsen) 6.2 mg kg1 soil. Prior to this study the
plant growth rates has led to such a plethora of field was under corn cultivation (1993).
research that it is now itself a specialised discipline of The climate of the region is classed as a thermic
plant physiology in its own right. The growth of a crop Mediterranean (FAO-UNESCO, 1973). Average rain-
passes through three consecutive phases which in their fall (for 30 years) is 1022 mm with 68% falling
entirety determine a quantitative relationship between between November and April, with an average air
production/yield and time. This relationship is temperature of 14.7 8C. Climatic data such as air
described by the sigmoidal growth curve. temperature, air humidity, global radiation, wind
The logistic equation (Hunt, 1982) velocity and precipitation were measured from an
a automatic weather station installed 40 m from the
W (1) experimental site. During the cultivation periods of
1 ebct
1994 and 1995, maximum air temperature was greater
is a mathematical expression describing the sigmoidal than 30 8C in JuneAugust, whilst the minimum
growth curve, i.e. the growth of a plant, where W is the temperature ranged from 13 to 19 8C and 11 to 18 8C
N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113 3

for 1994 and 1995, respectively, over the same period. broadcast over the soil surface of each plot before
No difference in rainfall distribution during the two being incorporated into the soil to a depth of 0.15
experimental periods was observed. Briefly rainfall 0.2 m by hand. Topping took place when 50% of the
was 98 mm in 1994 distributed as 23% in May, 39% in plants in each plot were at full bloom (75 and 76 DAT
July and 38% in August. In 1995, rainfall was 101 mm in 1994 and 1995, respectively) and to a height such
distributed as 24% in May, 35% in July and 41% in that 2224 leaves remained. All cultivation practices
August. Tobacco plants were irrigated 12 times in took place in strict accordance with the guidelines of
1994 and 10 times in 1995 with a total irrigation of the National Tobacco Institute of Greece (1996).
350 mm each cultivation period. The quantity of water On an approximately weekly basis, three plants
used for irrigation was equal to the maximum were randomly selected from each experimental plot
evapotranspiration rate (ETc) which was estimated from 34 DAT until 96 DAT (full maturation and seed
from the potential evapotranspiration (ETp) and the production) resulting in the sampling of 54 plants per
crop constant for tobacco (Kc). ETp was calculated sampling date. The length of the various stages of
using the minimum and maximum air temperatures, tobacco development and the number of samplings
duration of solar radiation, relative humidity and wind during that period (National Tobacco Institute of
speed, using the Penman Equation (1948) as modified Greece, 1996) are shown below:
by Frere (1979). The value of the constant Kc for
tobacco during the growing season was taken as 0.4  recovery (adaptation): 3035 days, 1 sampling;
for the recovery period, 0.70.8 for the phase of rapid  knee-high: 1015 days, 2 samplings;
growth and elongation, 0.91.0 for the stage of  budding (rapid growth and elongation): 1015 days,
flowering and topping and finally 0.750.85 for the 2 samplings;
maturation and seed formation phase (Doorenbos and  flowering and topping, beginning of harvest: 2535
Pruit, 1977). Irrigation was carried out when the days, 2 samplings;
moisture content at 0.3 m was less than 40% of the  maturation and seed formation: 2025 days, 2
available water moisture. Soil moisture was measured samplings.
using tensiometers installed in the field.
The experimental field comprised 18 plots Each plant was extracted from the soil by digging a
(150 m2) arranged in 3 rows of 6 plots with a distance trench around it 0.3 m2 by 0.4 m deep and removing it
of 1.5 m between rows and 1 m between plots. On 18th as a block. The roots were washed well to remove all
May 1994 and 19th May 1995, each plot was planted traces of soil and the plants then separated into leaves,
up with flue-cured tobacco plants (McN-944) in 15 stalks and roots. Individual parts were then washed to
rows, each row 10 m long, with a distance of 1 m remove all traces of soil. The corresponding parts of
between rows and 0.5 m between plants (20,000 plants the three plants sampled were combined to produce a
per ha). Half of the plot was used to monitor DMA and bulk sample for each plant part and each experimental
nutrient uptake, the other half being used for plot. For statistical purposes, there were thus 18 re-
collection and processing of mature tobacco leaves plicates. The fresh weight of the bulk sample plant
to determine crop yield. Fertilisation was carried out parts was recorded and the samples were then dried to
each year using quantities appropriate for maximum a constant weight in an oven at 70 8C whereon dry
yield and quality production of flue-cured tobacco. weight was then recorded. The parts were then ground
The total amounts of fertiliser used were 6 g N m2, to pass a 250 mesh sieve. A portion of the ground
12 g P m2, and 24 g K m2. Applications were made samples was incinerated at 550 8C and the ash diss-
23 days before transplanting using 4 g N m2, olved in concentrated nitric acid. This solution was
12 g P m2 and 17 g K m2 using calcium ammonium used for the determination of Ca, Mg, P and K. Co-
nitrate (26% N), ordinary superphosphate (21% P2O5) ncentrations of Ca and Mg were determined using
and potassium sulphate (50% K2O). One month after atomic absorption spectrometry with an acetylene
transplanting, an application of 2 g N m2 and N2O and acetyleneoxygen flame for Ca and Mg,
7 g K m2 was applied in the form of KNO3 (13% respectively. Potassium was determined using flame
N and 46% K2O). The fertiliser was uniformly 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 during the growth period follows a sigmoidal curve
determined using the Kjeldahl method. The analysis of (Figs. 15) which can be accurately described by the
plant parts took place in accordance with the methods logistic equation:
reported by Page (1982). a
The dry weight of leaves, stalks and roots was W
1 ebct
determined for each sampling date and expressed as
g m2. The amount of N, K, P, Ca, and Mg taken up where W is the accumulation of DML, DMS, DMR,
and distributed within plant parts was calculated by DMAe and DMP at a time t, a is the maximum value of
multiplying the dry weight by the corresponding DMA, b the initial DMA, c an accumulation constant,
elemental concentration and expressed as g m2. The t the time in days after transplanting. The values for a,
leaves of plants not sampled for elemental analysis b, and c were calculated using Rosenbrocks method
were harvested and dried in a bulk-curing furnace, (Machura and Mulawa, 1973) with the statistical soft-
appropriately treated and the tobacco yield calculated ware application Statistica for Windows (StatSoft,
per hectare. 1995). The correlation coefficient r between predicted
After data collection, statistical analyses were and observed values is very high (Table 1) and ensures
performed in the following order to: the statistical integrity of the curve.
Figs. 15 show the accumulation curves for whole
(a) examine the effect of year and date on DMA and plants and plant parts resulting from the application of
nutrient uptake and distribution in whole plant and the logistic equation to the data. A study of these
plant parts; curves gives us the following information.
(b) determine whether data for the 2 years could be
combined and analysed, using Bartletts x2 test for 3.1. Dry matter accumulation
homogeneity;
(c) examine the adaptation of our data using non- Dry matter accumulation in whole plants and plant
linear regression to a logistic equation and parts was relatively slow until 35 DAT, indicating the
estimation of the parameters a, b, and c; slow adaptation of the plants to their new environment.
(d) statistically evaluate predicted and measured Similar periods of adaptation have been reported by
values. Raper and McCants (1967), Atkinson et al. (1977),
and Mylonas and Pangos (1980) for flue-cured, Burley
3. Results and discussion and orient type tobaccos, respectively.
At maturation in flue-cured plants 848 g DM m2
The distribution of rainfall, as well as other factors accumulated distributed as 356 g m2 in leaves,
influencing tobacco growth, such as daily minimum 277.2 g m2 in stalks and 240.49 g m2 in roots
and maximum air temperatures, was satisfactory (Fig. 5) representing 42.3%, 33% and 28.5%,
in both years. The average annual production of respectively, of total DM. The sum of DM in plant
processed leaves was 3450 kg ha1 (380.3). parts is not equal to the total DMA as a result of
An examination of the data showed that experi- smoothing applied during the regression analysis.
mental year and sampling date had no effect on DMA In the period between 34 and 96 DAT, between
nor nutrient uptake and distribution. Bartletts x2 test transplant and maturation, there is rapid DMA in plant
showed that combining the data from both years was parts and whole plants (Figs. 1a, 2a, 3a, 4a and 5a).
acceptable. In addition, deviation in the transplant date This period differs between plant parts both in respect
in the two experimental years was almost non-existent to onset as well as to duration. In particular, in leaves
and hence for any given sampling date the plants were this period of rapid DMA is between 41 and 75 DAT
at the same stage of development in each year. In the (Fig. 1a) in which 93% of total DMA is achieved,
analysis that follows, all values given are the averages corresponding to 39.4% DMP. Similarly, Suzuki et al.
of the data for the 2 years combined.The DMA in (1970) report rapid changes in DMA in Burley tobacco
leaves (DML), stalks (DMS), roots (DMR), aerial leaves from 30 to 75 DAT. In stalks, there is a period of
parts (DMAe = DML + DMS) and whole plant (DMP) 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. tobacco plant. The maximum daily DMA differs
This period of rapid DMA occurs in roots from 48 to between plant parts both in value and in date of onset.
90 DAT (Fig. 3a) where 82% of DMR is achieved, The maximum DMA in leaves, stalks and roots is
corresponding to 11.8% of DMP. observed on 56, 70 and 69 DAT, at 89.5, 72.4 and
From transplant to 48 DAT (i.e. the first half of 60.2 g m2, respectively (Figs. 1b, 2b and 3b).
the period from transplant to maturation) DMA is The rate of increase in DMP rises until 62 DAT
86.6 g m2 in leaves, 18 g m2 in stalks and 3 g m2 when it reaches its maximum, after which it declines
in roots, representing 23%, 6.5% and 1.3% of DMA in to zero when DMA is at its maximum. Fig. 6 shows
leaves, stalks and roots or 9.0%, 2.0% and 0.4%, that the maximum growth rate, i.e. maximum daily
respectively, of DMP at maturation. DMA, is achieved at time t where the second factor of
DMA in leaves, stalks and roots per unit time is Eq. (3) tends to zero and DMA is at half its maximum.
calculated from Eq. (2) and expresses the rate of The same pattern in daily DMA is observed in plant
growth, i.e. the daily DMA in the various parts of the 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 where U is nutrient uptake (g m2) at time t, a is
appears towards the end of the budding stage (early maximum nutrient uptake (g m2), b the initial nutri-
floweringtopping stage). ent uptake (g m2), c a nutrient uptake constant, t time
in days after transplant and e the natural logarithm.
3.2. Nutrient uptake Again the sum of uptakes is not equal to the total
uptake due to smoothing in the regression analysis.
The uptake of nutrients in whole flue-cured tobacco
plants, as well as the uptake and distribution in 3.2.1. Nitrogen uptake
individual plant parts follows a sigmoidal curve The pattern of N uptake in whole plants and in plant
(Figs. 15) described by the logistic equation: parts is shown in Figs. 1a, 2a, 3a, 4a and 5a. A period
of rapid change in N uptake is observed between 41
a and 68 DAT where 85% of whole plant N is taken up.
U
1 ebct 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 respectively (Fig. 5a). During the growth period from
rapid change in uptake occurred between 41 and 68 transplant to 48 DAT (i.e. the first half of the growth-
DAT (Fig. 1a) where 93% of total leaf N was taken up, maturation phase) N taken up and distributed in
corresponding to 56% of whole plant N. The leaves, stalks and roots was 2.9, 0.5 and 0.13 g m2
maximum daily N uptake distributed in leaves was corresponding to 16%, 2.6% and 0.7% whole plant N,
found on 54 DAT at 2.7 g m2, in stalks 67 DAT at respectively. During the floweringtopping stage, the
0.86 g m2 and in roots 64 DAT at 1.1 g m2 (Figs. 1b, leaves, stalks and roots contained 10.7, 2.3 and
2b and 3b). The amount of N in the whole plant at 4.22 g N m2, respectively. These values represent
maturity was 18.24 g m2, distributed as 10.9 g m2 98%, 72% and 93% of total uptake in leaves, stalks
in leaves, 3.22 g m2 in stalks and 4.6 g m2 in roots, and roots, respectively, and correspond to 58%, 13%
representing 60%, 18% and 25% whole plant N, 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- DAT and in roots between 48 and 75 DAT (Figs. 1a, 2a
tion of small quantities of N to the leaves of flue-cured and 3a) where 93%, 92% and 93% of total leaf, stalk
tobacco after flowering is desirable to ensure good and root K was taken up, corresponding to 18%, 33%
quality. and 22.5% of whole plant K. The amount of K in the
whole plant at maturity was 18.54 g m2, distributed
3.2.2. Potassium uptake as 7.6 g m2 in leaves, 6.7 g m2 in stalks and
A period of rapid change in K uptake is observed 5.3 g m2 in roots, representing 41%, 36% and 29%
between 41 and 68 DAT where 81% of whole plant K whole plant K, respectively. These values represent
is taken up (Fig. 5a). The maximum daily uptake of K 98%, 62% and 93% of total K uptake in leaves, stalks
is observed on 61 DAT at 4.6 g m2 (Fig. 5b). In and roots, respectively, and correspond to 40.5%, 23%
leaves, the period of rapid change in uptake occurred and 27% of whole plant K uptake. During the growth
between 41 and 68 DAT, in stalks between 41 and 90 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 uted in stalks occurred 73 DAT at 1.86 g m2 and in
distributed in leaves, stalks and roots was 1.9, 0.7 roots 64 DAT at 1.33 g m2 (Fig. 1b)
and 0.14 g m2 corresponding to 10.3%, 3.9% and The quantities of N and K taken up and distributed
0.8% whole plant K, respectively. in aerial parts of the plant are approximately the same
The distribution of K in tobacco plants indicates at 18.5 g m2. McCants and Woltz (1967) report that
that K is essential even after topping. McCants and while flue-cured tobacco takes up greater quantities of
Woltz (1967) report that application of a K containing K than N, greater quantities of K-fertiliser must be
fertiliser 34 weeks after transplanting favours the added for the further reason that a high K uptake
production of good tobacco. The maximum daily K improves tobacco quality.
uptake distributed to leaves was found on 54 DAT at
1.91 g m2 (Fig. 1b) showing that K must be present in 3.2.3. Calcium uptake
significant quantities even at the early stages of A period of rapid change in Ca uptake is observed
development. The maximum daily K uptake distrib- 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 11

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, During the floweringtopping stage, the leaves, stalks
respectively. These values represent 95%, 70% and and roots contained 0.5, 0.24 and 0.24 g P m2,
90% of total uptake in leaves, stalks and roots, respectively. These values represent 98%, 96% and
respectively, and correspond to 48.16%, 16% and 26% 90% of total uptake in leaves, stalks and roots,
of whole plant Mg uptake. The maximum daily Mg respectively, and correspond to 49%, 25% and 26% of
uptake distributed in leaves was found on 56 DAT at whole plant P uptake. It is noticeable that P uptake
0.38 g m2, in stalks 67 DAT at 0.17 g m2 and in occurs during the first stages of development prior to
roots 64 DAT at 0.22 g m2 (Figs. 1b, 2b and 3b). topping. The maximum daily P uptake distributed in
leaves was found on 52 DAT at 0.12 g m2, in stalks
3.2.5. Phosphorous uptake 61 DAT at 0.06 g m2 and in roots 64 DAT at
A period of rapid change in P uptake is observed 0.07 g m2 (Figs. 1b, 2b and 3b).
between 41 and 75 DAT where 96% of whole plant P is
taken up (Fig. 5a). The maximum daily uptake of P is 3.2.6. Aerial plant parts
observed on 55 DAT at 0.24 g m2 (Fig. 5b). In leaves, The DMA in aerial parts of the plant exhibits a
the period of rapid change in uptake occurred between period of rapid change between 48 and 90 DAT
34 and 61 DAT, in stalks between 41 and 75 DAT and (Fig. 5a). Up to 48 DAT 12% DMA was produced and
in roots between 48 and 90 DAT (Figs. 1a, 2a and 3a) 18% N, 13.4% K, 17.6% Ca, 15.3% Mg and 17.2% P
where 94%, 95% and 99% of total leaf, stalk and root P of the whole plant content has been taken up and
was taken up, corresponding to 46%, 24% and 26% distributed in aerial parts. Atkinson et al. (1977),
of whole plant Mg. working with Burley tobacco, report that only 18%
The amount of P in the whole plant at maturity DMA of the aerial plant parts is produced during the
was 1.0 g m2, distributed as 0.5 g m2 in leaves, first half of the growth period. Grizzard et al. (1942),
0.25 g m2 in stalks and 0.27 g m2 in roots, working on flue-cured tobacco, report that during the
representing 49%, 25% and 27% of whole plant P, first half of a 90-day cultivation period, 18% of DMA
respectively. During the growth period from transplant was apportioned to the aerial parts of the plant.
to 48 DAT (i.e. the first half of the growth-maturation The maximum daily DMA in aerial parts of the
phase) P taken up and distributed in leaves, stalks and plants was observed 62 DAT at 159.9 g m2 (Fig. 4b).
roots was 0.16, 0.01 and 0.01 g m2 corresponding to The maximum daily uptake and distribution of N in
16%, 25% and 26% of whole plant P, respectively. 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 soil must have sufficient nutrients available to supply
in aerial parts was observed 57, 56, 58 and 55 DAT at the needs of the plant.
3.13, 2.98, 0.54 and 0.19 g m2, respectively (Fig. 4b). The greater part of N and Ca taken up by flue-cured
It is notable that the maximum daily uptake of all tobacco is distributed in leaves (60%) meaning that a
nutrients in aerial parts took place 57 days prior to the substantial proportion of these nutrients is removed at
attainment of maximum DMA. Raper and McCants harvest and should be replaced by appropriate
(1967) report that in flue-cured tobacco, 50% of N and fertilisation. In contrast, K, though taken up in equal
K of the plant is distributed to aerial parts during the quantities as N is distributed throughout the plant
early growth stage, specifically from 35 to 42 DAT and differently with 65% remaining in stalks and roots
that the uptake and distribution of Ca, Mg and P occurs after harvest.
at a relatively steady rate throughout the growth Hence for coarse-textured soils a split application
period. Atkinson et al. (1977), Bruns and McIntosh of 1/3 N and K fertiliser 23 days prior to transplanting
(1988) and Bertinuson et al. (1970) working with and 2/3 2530 DAT is recommended. For fine-textured
Burley, Maryland and shade-grown wrapper tobacco soils 2/3 prior to transplanting with an additional
respectively report that the daily uptake and distribu- application of the remaining 1/3 2530 DAT is
tion of nutrients to aerial parts was greater during the recommended in order to fulfil the plants require-
later stages of development. ments.
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 Acknowledgement
leaves, resulting in their removal from the field at
harvest. In contrast, the major portion of K taken up by The authors would like to thank Ms. Susan Coward
plants was distributed in stalks (36%) and roots (29%) for her help in the preparation of this paper.
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 References
leaves) and those remaining in the field after harvest
(roots and stalks). 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
4. Conclusions Connecticut shade-grown wrapper tobacco for three consecutive
years. Tob. Sci. 14, 155157.
During growth there is a period when DMA in plant Bruns, H.A., McIntosh, M.S., 1988. Growth rates and nutrients
parts occurs at an intense rate. The time of onset of this concentrations in Maryland tobacco. Tob. Sci. 32, 8287.
Ceotto, E., Castelli, F., 2002. Radiation-use efficiency in flue-cured
period and its duration vary with different plant parts.
tobacco (Nicotiana tabaccum L.): response to nitrogen supply,
Maximum daily DMA occurs when 50% of the climatic variability and sink limitation. Field Crops Res. 74,
maximum plant DMA has been achieved. 117130.
During growth there is a period of rapid change in Collins, W.K., Hawks, S.N., 1993. Principles of Flue-cured Produc-
nutrient uptake whose onset and duration vary not only tion. N. C. State University, Raleigh, NC.
with regard to the time of onset and duration but also to Doorenbos, J., Pruit, W.O., 1977. Crop water requirements. Irriga-
tion and Drainage. Paper 24 (revised). FAO, Rome.
specific nutrient, and plant part. Maximum daily Elliot, J.M., 1978. A survey of flue-cured tobacco grown in Ontario
nutrient uptake in aerial parts of the plant occurs in 1976. Part II. Nutrient elements. Lighter 47, 1921.
approximately 1 week prior to the maximum daily FAO-UNESCO, 1973. Carte bioclimatique de la zone Mediterranee.
DMA. O.N.M., Paris.
The period of rapid DMA and nutrient uptake in Flower, F.C., 1999. Field practices. In: Layten Davis, D., Nielsen,
M.T. (Eds.), Tobacco, Production, Chemistry and Technology.
flue-cured tobacco coincides with the knee-high and Blackwell Science, Oxford, UK.
budding (rapid qrowth and elongation) stage (between Frere, M., 1979. A Method for the Practical Application of the
41 and 75 DAT). Consequently during this period, the Penman Formula for the Estimation of Potential Evapotranspira-
N.K. Moustakas, H. Ntzanis / Field Crops Research 94 (2005) 113 13

tion and Evaporation from Free Water Surface. FAO, Rome, tobacco.. In: Proceedings of CORESTA Congress, 2024
AGP:Eco/1979/1. November, Manila, The Phillipines.
Hunt, R., 1982. Plant Growth Curves. Edward Arnold, London. National Tobacco Institute of Greece, 1996. Guidelines for Tobacco
Grizzard, A.L., Davies, H.R., Kangas, L.R., 1942. The time and Production (in Greek).
rate of nutrients absorption by flue-cured tobacco. Agron. J. 34, Page, A.L., 1982. Chemical and microbiological properties. In:
327339. Page, A.L. (Ed.), Methods of Soil Analysis. Part 2. Agronomy
Machura, M., Mulawa, A., 1973. Rosenbrock function minimization 9.. ASA, Inc., Madison, WI.
Algorithm 450. Commun. ACM 16, 482483. Penman, H.L., 1948. Natural evaporation from open water, bare
McCants, C.B., Woltz, W.G., 1967. Growth and mineral nutrition of soils s and grass. Proc. Roy. Soc. A 193, 120146.
tobacco. Adv. Agron. 19, 211265. Raper, C.D., McCants, C.B., 1967. Nutrient accumulation in Flue-
Miner, G.S., Tucker, M.R., 1990. Plant analysis as an aid in cured tobacco. Tob. Sci. 11, 190.
fertilizing tobacco. In: Westerman, R.L. (Ed.), Soil Testing Statsoft Inc., 1995. Statistica for Windows. tatSoft, Inc., Tulsa, OK.
and Plant Analysis SSSA Book, Ser. 3. SSSA, Madison, WI, Suzuki, M., Shinochara, T., Kona, K., 1970. Studies on the growth
pp. 645657. and the absorption of nutrients by Burley tobacco plants growth
Mylonas, V.A., Pangos, E.A., 1980. Dry matter and nutrient on Ando soils (Kuruboku). Morioka Lab. Tob. Exp. Sta. Bull.
accumulation in various plant parts of oriental neutral type 3, 91109.

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