Rev. Fac. Agron. (LUZ). 1997, 14: 507-516
Phenology and yield of maize for different sowing dates in a tropical
mountain environment
Efectos de diferentes fechas de siembra sobre la fenología y
producción de maíz en montañas tropicales
Recibido el 24-05-1996 l Aceptado
el 24-02-1997
1. Universidad de Los Andes. Facultad de Ciencias Forestales y Ambientales. Instituto de
Investigaciones Agropecuarias. Apartado 77 La Hechicera Mérida 5101, Venezuela.
Ramón E. Jaimez
Abstract
Experiments were carried out in order to examine the relationship
between leaf appearance and thermal time under field conditions and to improve a model
that simulates development and production of maize in tropical mountain environments. The
comparative phenology and productivity of the tropical highland maize cultivar Santa Rosa
were analized as a function of different sowing dates. The relationships between leaf
appearance rate and thermal time for both sowings dates tended to be basically curvilinear
and agreed with results shown in the literature. The number of leaves depended on daily
maximum and minimum temperature variations and on the stage of plant growth. These
affected the timing of anthesis and other development phases. The average yield and number
of ears was constant for both sowing dates
Key words: Zea mays, phenology, maize model, sowing dates.
Resumen
La variedad de maíz Santa Rosa fue sembrada en diferentes épocas de
siembra (enero y junio) a una altura de 1900 m con la finalidad de evaluar la aparición
de hojas en función del tiempo termal acumulado, el número de hojas y producción. La
relación entre el tiempo termal y la aparición de hojas en ambas épocas de siembra
obedece a una relación curvilínea, la cual se asemeja a otros reportes. El número de
hojas final va a depender de las variaciones de máximas y mínimas temperaturas en la
superficie del suelo en los primeros estadíos de crecimiento. La producción, el número
de mazorcas por planta y el número de granos por planta se mantuvieron constantes. Los
datos reportados ayudan a mejorar modelos de simulación de maíz en condiciones de
ambientes tropicales de altura.
Palabras claves: Zea mays, fenología, modelos de maíz, época de siembra.
Introduction
Maize (Zea mays L.) is the most widely grown cereal in Venezuela
producing more than 1.1 million tons over the last few years (6). Maize is mainly produced
in the lowland zones during the wet season, while in mountain areas (1500 m elevation), it
is planted during both the dry and the wet seasons.
Limited studies about dry matter production and phenology in the wet
season have been carried out in lowland regions of Venezuela (10, 13, 14, 8) and little
information exists on effects of planting dates on production and phenology. There are
also little data available for germplasm growing above 1600 m.a.s.l. under tropical field
conditions (1). According to studies in temperate zones, natural changes of temperature
and photoperiod have effects on phenology and production (19, 20, 16). Different results
were also found for tropical germplasm (7). In temperate regions yearly temperature
changes from season to season fluctuate from 8 to 12 °C while in tropical zones the
fluctuation is lower (from 3 to 4 °C). Due to this difference, the range of temperature
change effects on phenology and yield could vary between tropical and temperate maize.
In spite of adequate nutrients, water and weed control, the effects of
temperature on developmental events are difficult to evaluate because phenology is the
result of the interaction of other climatic factors. However, several authors have
indicated that leaf appearance rate and number of leaves are closely related to
temperature (16, 19, 20). They determined more precisely other phenological events such as
tassel and silking dates. It is often assumed that there is a linear relationship between
leaf appearance and day degrees, but this is not clear. Relationships between accumulated
temperature and leaf temperature rate have also been studied in tropical conditions (9,7)
and it is necessary that information on phenology and dry matter partitioning is obtained
to provide a framework for predicting and validating appropriate maize models for tropical
environments. These would lead to improved production.
The main objective of this study was to determine phenological periods
and leaf appearance for the Santa Rosa germplasm in a mountain tropical environment and
determine how they were influenced by sowing dates. A second objective was to obtain
knowledge about yield potential and to generate growth data to validate maize models.
Materials and methods
Experiments were conducted at the Universidad de Los Andes,
Agricultural Research Station (IIAP) in Venezuela (latitude 8° 38' N, longitude 71° 10'
W, altitude 1 900 m), during June 1991 through July 1992. The soil at this location is a
Humitropep Typic (12). Mean annual precipitation is 2 094 mm with a marked dry season from
December to March. The natural day length varies between 12.5 h and 11.5 h during the
year.
Cultural Practices: The Santa Rosa germplasm showing a high production
for mountain zones (1) was planted on June 27, 1991 and January 20, 1992. Two Kernels were
sown about 6 cm deep at 25 cm intervals and thinned to 40,000 plants ha-1. When
sowing, the plots were fertilized with 100 kg ha-1 of N, P and K each. Then,
additional fertilization was carried out using 100 kg ha-1 of N and K at 54
days after sowing (DAS). Plots were irrigated after 3 or 4 rain-free days. Diseases and
pests were controlled throughout the experiment. Both trials were replicated three times
in a completely randomized block design. Santa Rosa formed part of an experiment with
other genotypes of maize.
Measurements and data analysis: Emergence date was recorded when 50 %
of the plants had some part visible at the soil surface. At approximately 15 -17 DAS, 6
plants from each plot were selected and tagged for phenology measurements. From this time
to tasseling (50 % of plants with visi-ble tassel) each plant was measured twice a week
for last fully expanded leaf (when the ligule is visible). Silking date was recorded when
50 % of the plants had silks outside their husks. Physiological maturity was assumed to be
reached when in 50 % of the selected plants, 50 % of the grain possessed a black area (4).
Daily thermal time was calculated from maximum and minimum temperatures using 8 °C as the
base temperature (5).
Ears were removed and ear number, and potential grain number were
determined. Plants were oven dried at 75 °C for 7 days in order to determine dry weights
of leaves, stems, roots and grains. In addition, a Student's t test was used for
statistical analysis. In order to compare regressions between sowing dates an F test was
carried out (15)
Results and discussion
The natural day length for the June sowing was nearly 12.5 h during the
first 40 DAS. Daylength gradually decreased to 11.5 h at physiological maturity (158 DAS).
The January sowing daylength was approximately 11.5 h but increased later. At 47 DAS
daylength was 12 h and at physiological maturity (168 DAS) it was around 12.5 h (2, 3).
The average temperature was similar in both sowing dates throughout the life-cycle (17.8
°C, 17.3 °C in the June and January sowings respectively). However, for the June sowing
temperatures were higher than for the January sowing during the first 77 DAS (figure 1).
From this time until reaching maturity mean temperature of the June sowing decreased while
mean temperature for the January sowing increased. Radiation was similar in both
experiments (average 14.90 MJ/m2). Lower values were found to be related to
days-precipitation days. Soil surface minimun temperature was lower for the first eighty
days for the January sowing (figure 2).
Figure 1. Daily minimum and maximum temperatures during sowing on
June 20, 1991 and January 21, 1992.
Seedling emergence for the January sowing was reached within six days
after planting, while for June it occurred within five days. Corn cultivated in zones
above 1 700 m takes around thirty days more to reach physiological maturity than
cultivated corn in the lowlands plains (200 - 450 m) (8). Development was slower for the
January sowing compared to the June sowing (table 1). Anthesis of the January sowing
occurred at 721 ocd while for the June sowing it occurred a 705 ocd. This delay
was reflected in the greater number of leaves formed during January (table 2). Lower
minimum temperatures from sowing to tasseling are related to the greater number of leaves
formed. However, the January sowing experienced higher minimum and maximum temperatures
from tasseling to physiological maturity. This, however, did not accelerate the
phenological events. On the contrary, the duration of phenological stages was longer
(table 1).
Although temperature variations from cultivation to anthesis for both
planting periods are similar, this minimum variation apparently influences the number of
leaves. Average minimum and maximum temperatures for January were from 0.67 to 0.4 °C
lower with respect to average minimum and maximum temperatures for the June sowing. These
results fit those found by Warrington and Kanemasu (20) who, under controlled conditions,
found a curvilinear relationship between the number of total leaves and temperature. For
semi-tropical arid regions in Australia Muchow and
Figure 2. Soil temperature for first 80 days after sowing in June,
1991 and January, 1992.
Carberry (9) obtained a greater number of leaves for an August sowing
with respect to sowing carried out in October and February. In their experiment minimum
temperatures during the first 50 days of the August sowing were less than the minimum
temperatures of the October and February sowing. For this study 2.5 more leaves were
observed in the January sowing with a difference of 0.5 °C, while Muchow and Carberry (9)
showed a 1.7 leaf diffe-rence for the Delkalb XL82 hybrid between August and February with
a greater difference of temperature.
Table 1. Duration of phenological events (days and thermal time
(°Cd)) of cultivar Santa Rosa sown at two differents dates. Thermal time and days
estimated from emergence.
Sowing date |
Emergence |
50 % Tasseling |
50 % Silking |
Maturity |
|
°Cd |
days |
°Cd |
days |
°Cd |
days |
°Cd |
days |
Jun. |
44.4 |
5 |
705 |
70 |
788 |
78 |
1480 |
153 |
Jan. |
53.5 |
6 |
721 |
76 |
797 |
83 |
1601 |
161 |
Table 2. Rate of leaf appearance (leaves/day)during four phases of
leaf development and final leaf number for two sowing dates for cultivar Santa Rosa. Error
indicates standard deviation.
Sowing date |
Average rate of leaf appearance |
Final leaf number |
|
5-8 |
9-12 |
13-16 |
17-21 |
|
Jun. |
0.20 |
0.22 |
0.36 |
0.37 |
17.5 ± 0.5 |
Jan. |
0.21 |
0.23 |
0.34 |
0.41 |
21.4 ± 0.7 |
On the other hand, the relationship between the number of leaves and
the variation in temperature is a characteristic of the cultivar (17, 20), wich also
depends on the daily minimum and maximum temperature variations of each region. Lower soil
surface minimum temperatures registered during the first 45 DAS for the January sowing
days (figure 2) probably have a greater effect on the number of leaves. This coincides
with the temperature sensitive phase which influences the total number of leaves (17). For
the June sowing, soil surface temperatures were similar to the minimum air temperatures
(figure 1).
Leaf number and leaf appearance showed a curvilinear relationship with
thermal time (figure 3), consistent with findings by Muchow and Carberry (9) for the
semiarid tropical zone in Australia but not with findings by Manrique and Hodges (7) who
encountered a linear relationship in Hawaii for the hybrid Pioneer X304C. Until the
appearance of leaf number 12, for both trials, rates were maintained constant and did not
present significant differences. From the twelfth leaf, rates were around 38 % faster.
These variations are consistent with those found for hybrids in the field as well as
laboratory conditions in temperate zones (16, 18). However Tollenaar et al. (18)
found appearance rates to be 10 % lower from the third to the sixth leaf. Rates found for
this study's cultivar were higher than those reported by Warington and Kanemasu (19) and
Muchow and Carberry (9); that is, they were higher for the first twelve leaves as well as
for the apppearance rate after the twelfth leaf. Depending on minimum and maximum
temperature variation, the maximum appe-arance rate changes. Possibly such a variation
will be exaggerated as differences between minimum and maximum temperatures increase.
The relationship between thermal time and leaf appearance rate on the
cultivar has yet to be established. Equations which differentiate behaviors of leaf
appearance rate should form part of growth models and corn phenology in order to take in
account faster rates in the last formed leaves. F test showed no differences in linear or
exponential regressions, although the fitted exponential regression has a standard error
of estimate 10 times better for both sowings of Santa Rosa and all germplasms (table 3).
For the relationship between thermal time and number of fully expended leaves (NEFL) the
use of the exponential regression equation (NEFL = 1.16 exp (0.00245CTT), where CTT is
cumulative thermal time (table 3) is advisable. The incorporation of this equation could
improve the prediction of maize phenology for tropical areas. The equation found is
similar to that of Muchow and Camberry (9) for maize in semiarid tropical zones in
Australia.
Figure 3. Relation between expanded leaf number and accumulative
thermal time from emergence in June and January sowing. Each point is an average of 15
plants selected.
Biomass accumulation and yield. Significant differences were not
found in weight accumulation for expanded leaves, stalks (weight of the rest of the plant
minus expanded leaves and grains), and roots (data no showed). Leaf weight for the final
harvest was an exception although the differences were not significant for the two sowing
dates (table 4). The number of grains per ear was not significantly different for the two
sowing dates. A significant effect was not established between sowing time with
production, number of grains and harvest index (HI). (table 4). The average production of
both trials was 685 g/m2. This is similar to production rates of other corn
varieties grown at lower altitudes (between 60 - 500 m) where greater extensions were
utilized for cultivating corn and where production was between 400 - 694 g/m2 (10,13,14).
Although most of the sowing is done in the rainy season (April to October), the wide
production range may be attributed to the availability of soil water which depends on
unpredictable precipitation schedules. The number of ears of corn was relatively constant
for both crop periods (1.6 ears/plant). This gives approximately 581 potential grains per
plant. This value is high compared to other tropical hybrids (11, 8).
Table 4. Effect of sowing date on dry yield (g m-2),
grain number per ear and harvest index. (error indicates standard deviation). values with
same letter are not significantly different at P = 0.05 according to Student's test.
Variable |
Sowing date |
|
Jan. |
Jun. |
Grain-yield |
667 ± 105a |
611 ± 60.5a |
Grain number |
373 ± 46.5b |
401 ± 46.5b |
Harvest Index |
0.40c |
0.47c |
Table 3. Regression analysis for linear and exponential models for
June and January sowing of Santa Rosa germplasm.
Lineal model Y = a + bx |
Exponential model Y = exp (a + bx) |
Parameter |
intercept |
slope |
intercept |
slope |
Estimate |
-0.7445 |
0.0268 |
1.1616 |
2.565E-3 |
Standar error |
0.4131 |
8.175E-4 |
0.0587 |
1.162E-4 |
T value |
-1.8019 |
32.8498 |
19.76 |
22.065 |
Prob.level |
0.9671 |
0.0000 |
0.000 |
0.0000 |
R-squared |
98.90 |
97.59 |
|
|
Stand. error of estimate |
0.574 |
0.0816 |
|
|
F-ratio |
1079.10 |
486.86 |
|
|
Finally, the present work adds new information which may be included in
corn growth and development models adapted to the characteristics of tropical genotypes.
Furthermore, it demonstrates the need to develop a broader data base to elaborate
coefficients more in line with the phenological characteristics which tropical varieties
present.
Acknowledgements
The author gratefully acknowledges Lic. Fernando Castro, Eng. Ciro
Dávila and their group of workers for their assistance in the field work at the IIAP
station Santa Rosa. My appreciation also goes to researchers of the PAN EARTH project;
Venezuela case study. This work was supported in part by the Universidad de Los Andes
(CDCH-I328-90) and CONICIT (S1-2123)
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