Seasonal Variation of Biomass Partitioning and

Nutrient Use Efficiency of Corn (Zea mays L.)

Kate Mariel T. Colarina a,* Pompe C. Sta. Cruza
a
Institute of Crop Science, College of Agriculture and Food Sciences, University of the Philippines Los Baños, Laguna 4031, Philippines

A R T I C L E I N F O
A B S T R A C T
Keywords:
Seasonal variation Climate change is becoming inevitable and irreversible, and the Philippines is one of the most vulnerable nations
Biomass partitioning to its impacts. With temperature and precipitation as the major climatic factors affecting corn production, a field
Nutrient uptake study was conducted during the 2018 dry season and 2019 wet season in Ligao City, Albay and Lambunao, Iloilo
Nutrient use Efficiency to investigate the effects of seasonal variations on the BP and NUE of corn (Zea mays L.). Significant differences
Zea mays L. between the two seasons were noted for: 1) Biomass partitioning to a) kernel, b) stem, and c) tassel, husk, silk,
and cob; (2) Stover NPK uptake; (3) Kernel PK uptake; 4) Stover and kernel recovery efficiency of NPK; 5)
Internal utilization efficiency of PK; and 6) Physiological efficiency of K. Higher stover biomass, NU, majority of
the NUE parameters, and yield were observed under the wet season conditions, while higher KE biomass and IUE
were observed under the dry season conditions. Further studies on the sufficiency and toxicity ranges, for NU to
sustainably increase grain yield without risking environmental degradation and overcome the more intense
environmental conditions imminent with climate change, is recommended.

1. Introduction
projected a virtually certain increase in unusually warm days and nights,
Climate change is becoming inevitable and irreversible (IPCC, and a virtually likely increase in warm spells over most land areas. In
2021), and the Philippines is one of the most vulnerable nations to terms of precipitation, a likely increase in the frequency and intensity of
its impacts (Zhai & Zhuang, 2009; Cinco et al., 2014; Bathiany, 2018; rainfall is projected in high latitudes and in tropical regions.
Anderson et al, 2020). Studies have predicted the significant adverse Increases in temperature are expected to reduce corn yields (Bassu et
effects of climate change to agriculture and food security (IPCC, al., 2014; Anderson et al., 2020), especially in vulnerable areas, which
2007; UNCCS, 2019; Anderson et al., 2020; Fawzy et al., 2020; IPCC, accounts for about 43% of maize production (Pugh et al., 2016).
2022); and true to this, yellow corn production in the Philippines is Temperature increases to extreme values induce abiotic stresses such as
becoming more stagnant with only a 2.38% average increase from heat and drought stress which exude interactive effects leading to
2016-2020, as compared to the average of 5.23% increase from reduced photosynthetic, nutrient uptake and yield attributes in maize
2011-2015 (Philippine Statistics Authority, 2015, 2018, & 2021) (Hussain et al., 2019), inducing more significant yield reductions as
despite the ever-increasing use of fertilizers and hybrid seeds compared to the individual effects of these stresses (Zandalinas, 2018).
(Salazar et al., 2021). According to Ray et al. (2015), a third of observed crop yield variability is
Temperature and precipitation are among the major climatic explained by variability in climate.
factors affecting crop production (Easterling et al., 2007; From the analysis of Lesk et al. (2016), the global cereal production
Sathyaseelan et al., 2013; Bassu et al., 2014; Thornton et al., 2014; has decreased by 9-10% during 1964-2007 due to extreme heat and
Anderson et al., 2020; IPCC, 2022), more so for Philippine corn which drought. Studies by Brown and Rosenberg (1999) and Challinor et al.
is mostly rainfed and grown year-round. Salazar et al. (2021) have (2014) noted that increased temperature affects early maturity causing
noted in their Issues Paper on Corn Industry in the Philippines that less biomass accumulation and yield. For corn yields specifically, Zhao et
“the difference in weather patterns during the wet and dry season is al. (2017) have observed that a considerable reduction of 7.4% in corn
an important factor in appreciating the potential, limitation, and yields will be realized, without mitigation and adaptive strategies for
opportunities between Luzon, Visayas, and Mindanao”. climate change. Increasing temperature increases maize yield up to an
Defined as the shift in climate patterns caused by greenhouse gas optimum temperature of 29°C (Schlenker & Robers, 2009) to 30°C
emissions (Fawzy et al., 2020), Climate Change is affecting the world in a (Schauberger et al., 2017), but declines with further temperature
myriad of ways. Several studies have predicted the significant adverse increase as corn plants are sensitive to heat stress. Similar results have
impacts of climate change to agriculture and food security (IPCC, 2007; been reported by Lobell and Field (2007), wherein an 8.3% decrease in
UNCCS, 2019; Anderson et al., 2020; Fawzy et al., 2020; IPCC, 2022), as corn yield was observed for every 1°C rise from the optimum
agricultural yield is influenced heavily by abiotic stressors caused by the temperature. For every degree day spent above 30°C, Lobell et al. (2011)
severity and variability in climatic factors (Thornton et al., 2014) such as has reported a 1.7% reduction in corn yields under drought conditions.
rainfall, temperature, and atmospheric CO2, among others (Raza, 2019). With drought stress, a 40% water reduction rate causes a 39.3%
These adverse impacts will be felt to a greater degree by the developing reduction in corn yield (Daryanto et al., 2016).
and low-latitude countries (Zhai & Zhuang, 2009; Bathiany, 2018; Contributory to yield, biomass partitioning to the ear or the
Anderson et al., 2020) as such are already enduring intense climatic establishment of grain sink, is also largely affected by increased
factors (Mendelsohn, 2008), like the Philippines. temperature (Lafitte & Edmeades, 1997). The rates of dry matter
According to Sathyaseelan et al. (2013), increased temperature and partitioning and harvest index were reportedly impaired under heat
changed precipitation conditions are among the major climatic factors stress causing grain yield reductions (Rattalino Edreira & Otegui, 2012).
affecting crop production. The Intergovernmental Panel on Climate Suwa et al. (2010) have observed a larger decrease in the amount of 13C
Change (IPCC) (2022) has noted the significant increase in the partitioning to the ears more than the other plant parts as affected by
frequency and intensity of high temperatures and heavy precipitation, high temperature. Therefore, although plant biomass production
which are among the climatic-impact drivers. Temperature-wise, IPCC increased, grain yield is reduced due to the reduction in sink strength,
(2012) has

* Corresponding author.
E-mail address: k t c o l a r i n a 1 @ u p . e d u . p h ( K.T. Colarina)
K.T. Colarina and P.C. Sta. Cruz
rather than the source activity (Suwa et al., 2020; Chukwudi et al., numerical form of 4,745.27 kg ha-1 (Kim, Chemere & Sung, 2019), to cite
2021). Similar results were found by Lizaso et al. (2018) as stem a few percentage losses versus the normal climate.
biomass represented a higher percentage of plant biomass during Both intra- and inter-seasonal changes in precipitation also influence
hotter conditions. Results from the study of Rattalino Edreira and cereal yields (Rowhani et al., 2011). Strong positive correlation between
Otegui (2013) says otherwise, as heat stress has severe effects on both heavy precipitation and yield observed in July, versus a weaker
plant and ear growth rates but had less effects on the biomass correlation in August (Revadekar and Preethi, 2011), shows the
partitioning to the ear. In terms of drought, assimilates are translocated dependence of yield correlations on the month when the heavy
to the roots to improve water uptake, decreasing the shoot biomass precipitation occurred. Early-season moisture has also been reported by
(Leport et al., 2006; Eziz et al., 2017). Urban et al. (2015) to negatively affect crop yields by 2%.
Drought and heat stresses are also well known to decrease the Internally, yield is also correlated with precipitation during growth
nutrient uptake and photosynthetic efficiency of plants (Lamaoui et al., stages. Mtongori et al. (2015) noted the differing yield reductions by
2018), although accurate predictions are improbable due to the increased rainfall intensity during growth stages, where high
involvement of various interconnected metabolic processes (Ullah et al., precipitation during the vegetative stage resulted to a higher (5%)
2019). However, generally, nitrogen (N) uptake is most likely to decrease in yield, as opposed to the reproductive stages (2%). Similar to
increase, phosphorus (P) uptake to decrease, and potassium (K) uptake this is the result of the analysis of Rashid and Rasul (2011), where certain
to remain unaffected during drought stress (Fahad et al., 2017). amounts of rainfall increase yield at certain growth stages, specifically:
Amouzou et al. (2019) reported that both N and P uptake by maize 200-300mm for vegetative, and 200-250mm for reproductive. Moreover,
decreased under projected climate change scenarios. Decline was also Qi and Pan (2022) also reported the limiting effect of waterlogging during
observed in the partial factor productivity and internal nutrient use the early growing stages of maize.
efficiency of N by 10-47% and 5-33%, respectively for corn. Wang et al. As yield is affected by waterlogging due to high precipitation events, a
(2020) also studied the nitrogen use efficiencies of corn under drought certain impact is to be expected to biomass partitioning of corn. Hu et al.
stresses, and the resulting agronomic efficiency and partial factor (2022) noted that waterlogging during the third leaf stage (V3) reduces
productivity of N decreased by 27.4% and 25.8% respectively during carbon partitioned to corn ears (-53.1%), shanks (-46.5%), and nodes (-
the dry year as compared to the wet year. According to Bassirirad 71.5%), but increases carbon partitioned to leaves (9.6%) and tassels
(2000), heat stress-induced uptake reductions may be due to reduced (43.9%) compared to normal precipitation levels, leading to yield losses.
root mass and nutrient uptake per root area. Ultimately, the decrease in Similar to this is the study of Ren et al. (2014), wherein at the V3 stage,
corn production during heat and drought stress affects nutrient cycling, waterlogging affects the biomass partitioning of maize greatly, decreasing
uptake, and availability by adversely affecting various crop the distribution proportion of grain while increasing the proportions of
physiological functions (Fahad et al., 2017). stem and leaves. In terms of the root/shoot ratio, Ren et al. (2016) has
Challinor et al. (2014) found little evidence as to the potential of observed the greater influence of waterlogging on below- versus above-
maize to avoid yield loss in tropical regions. As mid and high latitudes ground biomass. In contrast, the study of Walne and Reddy (2021)
experience high temperatures optimal for corn growing, those which showed that the root/shoot ratio is unaffected, even when the weights of
are previously unsuitable for corn production will be able to produce the individual plant parts decrease with waterlogging.
more (Anderson et al., 2020); but areas that are currently suitable for As waterlogging caused by heavy precipitation promotes soil nutrient
corn production in the lower latitudes and tropical countries will reach losses, particularly runoff and leaching, nutrient uptake and use
extremes in the future, making them unsuitable for corn production. efficiency by plants is reduced (Kaur et al., 2019). Waterlogging also
According to the Assessment Report of the IPCC (2007), crop inhibits nutrient accumulation and distribution further causing reduced
productivity is expected to rise at mid- to high-latitudes as temperature nutrient use efficiency, leading to decreased yields (Ren et al., 2016, Ren
increases up to 1-3°C, while declines at lower latitudes for temperature et al., 2017).
increases up to 1-2°C. This is similar to the conclusion of Parry et al. Several studies have noted the effect of waterlogging to nutrient
(2005), as well as Rosenzweig et al. (2013), wherein climate impacts uptake and efficiencies. The study of Ren et al. (2017), showed that the
are more negative in the tropics and lower latitudes than in the mid- V3 stage of maize is most susceptible to waterlogging, significantly
and high-latitudes. Comparable findings were reported by the IPCC decreasing its total N accumulation, compared to control treatments.
(2014) in addition to indicating that the balance of impacts in the Similar results were obtained by Qi and Pan (2022) and Kaur et al.
higher latitudes, whether negative or positive, is still unknown. Such is (2018), wherein N uptake/accumulation, N use efficiency, and N partial
recommended by Rosenzweig et al. (2013), that further studies should factor productivity are decreased during waterlogged events.
be done to investigate the sustainability of agricultural production in Given the more intense climatic conditions associated with climate
these locations. However, as noted by Challinor et al. (2014), losses in change, this study aims to analyze the corn production of Albay and Iloilo,
aggregate production in cereal crops is expected not just for tropical as affected by temperature and precipitation. The main research
regions, but also for temperate regions as temperature increases by 2°C. questions posed are: 1) how are the yield and yield attributes
It is projected that climate change may cause an increase in the (comprising biomass partitioning, nutrient uptake, and nutrient use
frequency of heavy precipitation, particularly in high latitudes and efficiency) of corn affected by the contrast in climatic variables of the
tropical regions (IPCC, 2012), negatively affecting crop yields Philippine seasons?, and 2) what can be learned from this analysis to
(Easterling et al., 2007; Thornton et al., 2014). However, the effect of minimize the effect of climate change to sustain or even improve corn
heavy precipitation has a weaker association with yield than production?
temperature extremes (Vogel et al., 2019). Consequently, the IPCC
(2007) reported otherwise, as damages to the agricultural sector 2. Methods
caused by heavy precipitation were greater than those caused by
changes in temperature. 2.1. Time and Place of Study
Whether heavy precipitation affects yield more than temperature or
not, several studies still note the strong negative effects of high The experimental sites were selected based on their production
precipitation on crop yields, brought about by impacts on the plant performance for corn in the last 5 years (2012-2017), with the
physiological processes, physical damages, and delay in key field assumption that the recommendations which are proven effective for
operations (Revadekar & Preethi, 2011; Van der Velde, 2011; Kanchebe these sites could be repeated to other major corn-growing areas in the
Derbile & Abudu Kasei, 2012; Sun et al., 2020), due to waterlogging and Philippines. Due to the insufficient data from alternative areas, only two
flooding (Thornton et al., 2014; 2016; Anderson et al., 2020). Maize sites were selected to analyze the seasonal variations in biomass
yield losses from heavy precipitation have been reported to decrease to partitioning and nutrient use efficiency of corn, which are the localities of
around 13.3% (Song et al., 2020); 17% (Li et al., 2019); and an exact
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K.T. Colarina and P.C. Sta. Cruz
Ligao City, Albay and Lambunao, Iloilo (Figure 2). The 2018 dry season 3. Results
for both sites started from November 2018 to March 2019, while the
2019 wet season for Albay extended from July to November 2019, and 3.1. Agrometeorological data
for Iloilo from September 2019 to January 2020.
3.1.1. Temperature
2.2. Description of Experimental Sites
The maximum, minimum, and mean daily temperatures from November
2018 to January 2020 were obtained from two weather stations nearest to
Based on the modified Coronas Classification, the experimental sites
the places of study, which are Ligao City, Albay and Lambunao, Iloilo. From
in Ligao City, Albay experiences the Type IV climate or an evenly
these data, the average maximum, minimum, and mean temperatures were
distributed rainfall throughout the year, without a dry season. On the obtained for each month of the dry and wet seasons to analyze the effect of
contrary, Lambunao, Iloilo experiences the Type III climate, which is temperature on the yield and yield attributes of corn in the study areas.
characterized by no pronounced wet season, and a short dry season Albay and Iloilo both demonstrated lower average mean temperatures
occurring from December to February or from March to May. during the dry season (27.4°C and 27.9°C, respectively) than in the wet
season (28.7°C and 28.5°C, respectively) (Figure 1). In the Philippines,
2.3. Experimental Treatment and Design cooler temperatures may be observed among months close to January, and
warmer temperatures among months close to May; hence, cool dry seasons
occur in the months of December to February (PAGASA, 2014). This
The experiment was laid out in Completely Randomized Design
coincides with the 2018 dry season for both Albay and Iloilo, which
(CRD) with three replications and three farms per site. The unit plot explains the lower mean averages in the temperature of the dry season of
size for each treatment was 225 m2 (15 m x 15 m) with a 1 m distance corn production in both sites.
between each treatment plot. The widely-grown variety depending on
season per site was planted. During the dry season, Asian Hybrid J505 3.1.2. Rainfall
was used in Albay and Dekalb 6919S Genuity ® was used in Iloilo. On
the other hand, during the wet season, B9540G was used in Albay and The daily rainfall depth from November 2018 to January 2020 was
P3774R was used in Iloilo. obtained from two weather stations nearest to the places of study, which
The study consisted of two treatments which were: 1) site-specific are Ligao City, Albay and Lambunao Iloilo. From these data, the average
nutrient management (SSNM) and 2) control. The same management daily rainfall depth and the cumulative rainfall depth were obtained in
practices were applied to each treatment with the only difference being order to analyze the effect of rainfall to the yield and yield attributes of
corn in the study areas.
the rate of fertilizer application. SSNM treatment was based on the
Figure 1 shows the monthly variations in the rainfall parameters in
identified recommended fertilization rates specific to each area and Legazpi, Albay. During the 2018 dry season of corn production, the rainfall
season, which came out to be the same for both sites, but different depth averaged 8.33 mm daily, with a cumulative depth of 1190.40 mm for
depending on the seasons. During the dry season, the fertilizer rate was the entire season. These amounts were higher than the wet season average
160-70-170 kg N-P2O5-K2O ha-1, and 140-60-150 kg N-P2O5-K2O ha-1 daily and cumulative rainfall depths, which were 5.67 mm and 735.90 mm,
during the wet season. The control treatment was the checkplot respectively. The higher rainfall parameters observed in the region may be
without any fertilizer applied. attributed to the outlying data observed during December 2018; during
Land was prepared after soil samples were obtained from each which, the corn was at its early vegetative growth stage.
In Iloilo, the rainfall amounts are lower for the dry season as compared with
experimental site. Plowing followed by harrowing and leveling was
the wet season. The daily rainfall depth averaged 2.20 mm and 13.55 mm,
done to over-turn the soil and break the soil clods into smaller masses. respectively; and the cumulative rainfall were 203.50 mm and 1142.60 mm,
Dibbling method was used to sow the corn seeds by digging holes 20 cm respectively. Contrary to the Albay climatological data, the rainfall depth for wet
apart for each row. Two seeds were sown per hill, which were thinned season corn production in Iloilo is greater than its dry season corn production, as
no outlying data was observed.
down after 2-3 weeks to maintain 1 plant per hill. Plant spacing was 20
cm between hills in a row, by 75 cm between rows. The fertilizer
(a)
application for Site-Specific Nutrient Management (SSNM) was done in
two splits, i. e. basal application before planting and side-dressing at 28
days after sowing (DAS). Harvesting was done at physiological
maturity. December to February or from March to May.

2.4. Statistical Analysis

A series of tests were conducted for comparative and regression

analyses. This study used a 95% level of confidence with alpha = 0.05
for the hypothesis testing. The decision rule of the hypothesis testing is
that the null hypothesis was rejected when the p-value was less than
alpha = 0.05. Comparative analysis was used to assess if there are
significant differences between the considered variables. The following
tests were performed: 1) Welch F-test was used to test the hypothesis
of the data which satisfied the assumption of normality; 2) Mann-
Whitney U-test was used to test the hypothesis of the data which did
not satisfy the assumption of normality. Multiple regression analysis
was also done through the Pearson’s correlation coefficient to
determine if there is a significant correlation between the considered
variables.

10
K.T. Colarina and P.C. Sta. Cruz
with ranges of 1.86-4.13% for the dry season, and 1.46-3.89% for the wet
(b)
season, denoting an average organic carbon content in either season. The
NPK content of the soil was also initially analyzed, wherein both the total N
content and K content of the soils were high and similar for both seasons.
Moreover, varying values are noted for the P content with wide ranges of
2.5-36.0 ppm during the dry season, and 3.0-90.0 ppm during the wet
season, in which the increased amount may be attributed to the soil water
movement of P brought about by higher precipitation (Sawyer et al., 2000).

3.2. Effects of Cropping Season on Biomass Partitioning

The biomass partitioning of corn was obtained by collecting the

individual weights of the stover parts and grains. The aboveground
biomass of corn was disaggregated into four main components: 1) leaves
and leaf sheath (LLS); 2) tassel, husk, silk, and cob (THSC); 3) stem; and
(c) 4) kernel. During the dry season, significantly larger proportions of
biomass are allocated in the economic parts of corn, which are the THSC
and kernel, while the LLS and stem have higher mean weights during the
wet season (Fig. 2). As compared with the wet season, The percentage of
THSC and kernel components on the total corn biomass were higher by
4.9% and 11.4%, respectively; while LLS and stem were lower by 6.1%
and 10.2%, respectively, during the dry season.

(d)

Fig. 2. Absolute and relative biomass partitioning in corn at physiological

maturity during dry and wet seasons.

3.3. Effects of Cropping Season on Nutrient Uptake

In the absolute comparison between seasons (Fig. 3), corn plants

Fig. 1. Climatic variables vis-à -vis growth stages during (a) Albay dry season, grown in SSNM treatments had higher nutrient (NPK) uptake during the
(b) Albay wet season, (c) Iloilo dry season, and (d) Iloilo wet season. wet season cropping. Comparative analysis between seasons using the
Mann-Whitney U test indicates that all uptakes except for the N uptake in
With these figures, the following observations are made: 1) during vegetative
the kernel significantly differed between seasons. Kernel P and stover NP
growth stage, a) excessive rainfall on Albay DS, Albay WS, and Iloilo WS, and b)
sufficient rainfall on Iloilo DS; 2) during the reproductive growth stage, uptakes were significantly higher during the wet season, while kernel and
insufficient rainfall for both sites and seasons (Albay DS & WS, Iloilo DS & WS); stover K uptakes were significantly higher during the dry season.
and 3) during the maturity stage, a) insufficient rainfall on Albay DS and Iloilo
DS, and b) excessive rainfall on Albay WS and Iloilo WS.

3.1.3. Soil Properties

Initial soil analysis was done prior to the conduct of the experiment in
order to determine the key properties of the soils in the experimental set-
up. With this, the seasonal differences in the soil chemical properties,
including soil pH value, total organic matter (OM), and nitrogen,
phosphorus, and potassium (NPK) content of the Ligao City, Albay and
Lambunao, Iloilo soils were analyzed.
The pH of soil in both the study sites during the dry season is 5.4-6.2,
which is within the optimum range for corn (5.6-8.1) as reported in
several studies (Gale, Koenig & Barnhill, 2001; Albrecht, Ketterings &
Beckman, 2005; Setiawati & Yusuf, 2020). On the contrary, the pH of soil
during the wet season in Albay and Iloilo is 4.7-5.7, which suggests a
more acidic environment during the wet season corn production. This
Fig. 3. Absolute and relative nutrient (NPK) uptake by corn stover and kernel at
observed higher acidity may be due to: 1) leaching of soluble salts (e.g.
physiological maturity during dry and wet seasons.
Na+, CO32-, HCO3-) during increased precipitation (Jia et al., 2021);
and/or 2) sulfur dioxide (SO¬2) emission of the Mayon Volcano
distributed to the soil through acid rain deposition (Mouginis-Mark, Comparing Figure 3 with the seasonal variation in biomass
2005; PHIVOLCS, 2023). partitioning under the SSNM treatment, lower absolute kernel and stover
The total percent organic matter was comparable between seasons, nutrient uptakes were observed during the dry season as compared with
10
K.T. Colarina and P.C. Sta. Cruz
the wet season. Similarly, lower stover biomass was observed during in the kernel biomass partitioned during the dry season. That is,
the dry season; and on the contrary, higher kernel biomass was increased kernel N and P ensues increased kernel biomass.
observed during the dry season. Comparing the relative nutrient
uptakes brings consistent results where higher kernel and lower stover 3.5. Effects of Cropping Season on Nutrient Use Efficiency
uptake and biomass were observed during the dry season as compared
with the wet season. The study used five measures of nutrient use efficiency
analyzing crop productivity (Partial Factor Productivity and
3.4. Relationship between Biomass Partitioning and Nutrient Uptake Agronomic Efficiency), nutrient uptake (Recovery Efficiency),
and ability to transform nutrients into economic yield (Internal
Stover biomass (y) corresponded significantly (p<0.05) with the Utilization Efficiency and Physiological Efficiency), as affected by
nutrient (P and K) uptake (x), and revealed a linear relationship as the Philippine cropping seasons. Two of these measures (Partial
shown by the graph in Figure 4. Factor Productivity and Agronomic Efficiency) provided
insignificant results (higher values observed during the wet
season) on the comparison of efficiencies between seasons.

3.5.1. Apparent Recovery Efficiency (RE)

Generally, recovery efficiencies were relatively higher during the wet

season. Figure 6 shows that in the stover, the largest difference was
observed in K uptake, wherein the K recovery efficiency during the wet
season is 775% higher than the dry season; followed by N with 440%
increase, and P with 300%. On the other hand, in the kernel, the largest
difference was observed in P uptake, with 500% higher P recovery
efficiency during the wet season than the dry season; followed by K with
400% difference, and N with only 157.14%.

Fig. 4. Stover biomass partitioning and nutrient (PK) uptake at physiological

maturity in corn during the dry season.

Under the SSNM treatment, the linear equation BP = 127.10 SUP +

356.78 SUK – 323.15 was derived from the P and K uptakes and stover
biomass. The kernel uptake for N was not included in the equation since
it was not significant at p<0.05. The obtained equation signifies that the
changes in P and K nutrient uptake would result in corresponding
linear changes in the stover biomass partitioned during the dry season.
That is, increased P and K NU in the stover would result in increased
biomass partitioning to the stover.
In terms of the kernel nutrient uptake and biomass partitioning Fig. 6. Recovery efficiency of stover and kernel due to nitrogen, phosphorus,
(Table 8), there is a significant correspondence (p<0.05) of the kernel and potassium applications during dry and wet seasons. Vertical bars indicate
standard deviations.
biomass (y) with the nutrient (N and P) uptake (x), which revealed a
linear relationship as shown by the graph in Figure 5.
The comparative analysis resulted to significant values at p-
value<0.05, which indicates that: 1) seasons significantly affected the
NPK stover and kernel recovery efficiencies; and 2) the recovery
efficiency of corn is significantly greater during the wet season cropping.

3.5.2. Internal Utilization Efficiency (IE)

The internal utilization efficiencies resulted to contrasting values with

the PFP and AE, in which higher efficiencies during the dry season than
the wet season were observed (Fig. 7). The percentage differences of N, P,
and K are 7.57%, 113.92%, and 36.67%, respectively. The higher values
obtained during the dry season as compared with the previous increases
in the wet season for other nutrient use efficiency measures signifies that
the high amount of nutrients the corn plants were able to take up during
the wet season were not translated into economic yield.
Fig. 5. Kernel biomass partitioning and nutrient (NP) uptake at physiological
maturity in corn during the dry season.

Under the SSNM treatment, the linear equation BP = 97.60

KUN + 49.05 KUP – 163.52 was derived from the N and P
uptakes and kernel biomass. The K kernel uptake was not
included in the equation since it was not significant at p<0.05.
The obtained equation indicates that the changes in kernel
nutrient uptake would result in corresponding linear changes

10
K.T. Colarina and P.C. Sta. Cruz
partitioning. The regression analysis shows no significant
relationship between the recovery efficiency of nutrients in the
stover with the stover biomass partitioning of corn during the
dry season. This indicates that the changes in stover recovery
efficiency would not result in any changes in the stover biomass
partitioning during the dry season. On the other hand, in terms
of the kernel recovery efficiency, a significant relationship
between the recovery efficiency of N in the stover with the
stover biomass partitioning during the dry season was observed.
Under the SSNM treatment (balanced fertilization), the linear
equation BP = 311.94 REN + 67.55 was derived from the N
recovery efficiency and kernel biomass. The P and K recovery
efficiencies were not included in the equation since these were
not significant at p<0.05. The obtained equation in the study
Fig. 7. Internal Utilization Efficiency of Zea mays L. between seasons. Vertical indicates that the changes in kernel N recovery efficiency would
bars indicate standard deviations. result in corresponding linear changes in the kernel biomass
partitioned during the dry season, considering that N is the most
The comparative analysis resulted to significant values of P and K critical nutrient in corn production for optimum grain yield
efficiencies at p-value<0.05, which indicates that seasons have a (Miao et al., 2006).
significant effect on the IE. In terms of the internal utilization efficiency
of N, other biotic and abiotic factors may affect it other than the climate.

3.5.3. Physiological Efficiency 3.7. Effects of Cropping Season on Yield

The physiological efficiency of corn (PE) was analyzed to determine Results of the study reveals that between seasons, the control
the ability of the corn plants to transform fertilizer applied into treatment yield during the dry season was higher than the wet
economic yield. the N and K physiological efficiency values are season, while the SSNM treatment yield was higher during the wet
consistent with the internal utilization efficiency wherein these are season (Figure 9). As compared with the wet season, the dry
higher during the dry season. The P physiological efficiency values are season yield is 22.1% higher without fertilizer applied and is 7.3%
similar to the partial factor productivity and agronomic efficiency, lower with fertilizer applied.
which are higher during the wet season. The percentage differences of
N, P, and K are 14.83 %, 46.37%, and 33.94%, respectively. The
obtained values in the present study suggest that the plants were able
to take up a high amount of nutrients from the fertilizer applied, but as
reflected in the physiological efficiency, these were not translated into
economic yield.

Fig. 9. Grain yield of corn as affected by fertilizer treatments during dry and wet
seasons. Vertical bars indicate standard deviations..

The yield was not significantly different (P≤0.05) between

locations, and as a result, means were combined across sites. Using
the Welch F-test, no significant difference in grain yields between
seasons was observed. This may be true statistically, although
under field/farm level, a 7-20% difference in yield could be
Fig. 8. Physiological efficiency of Zea mays L. between seasons. Vertical bars relatively substantial. The non-significant difference in yields
indicate standard deviations. among seasons, with the contrasting significant difference in yield
attributes pique interest. However, the correlation of the various
Comparative analysis resulted to significant potassium efficiency at yield attributes to yield resulted in inconsistent outcomes due to
p-value<0.05, which indicates that seasons have a significant effect on the limited sample size.
the potassium PE or the ability of plants to transform potassium The difference between treatments was also analyzed where
fertilizer inputs into economic yield. In terms of the physiological both seasons have a significant difference between their control
efficiency of N and P, other biotic and abiotic factors may affect these and SSNM treatments, which indicates that fertilizer application
besides the climate. affects the yield production of corn.

3.6. Relationship between Biomass Partitioning and Nutrient Use 4. Discussion

Efficiency
4.1. Heat Stress, Excessive Rainfall, and Ear Biomass
As recovery efficiency is the only measure that was divided
into its stover and kernel NUE, this was used in finding out the Biomass partitioning is significantly different between seasons, except
relationship between nutrient use efficiency and biomass for the leaves and leaf sheath part. This suggests that the climatic

10
 K.T. Colarina and P.C. Sta. Cruz
conditions relative to the dry and wet seasons affect the allocation of yield of corn would be possible by investigating its nutrient uptake during
biomass of corn. From the agrometeorological data obtained, the average the growth seasons.
mean temperature, maximum temperature, daily rainfall depth, Applied fertilizers to the soil surface are subject to various reactions
cumulative rainfall depth, and pH of dry season corn are correspondingly which may limit their availability to the corn plant, lowering its uptake. In
lower than the wet season. terms of N and P, higher uptakes were observed during the wet season, as
The lower kernel mean weights during the wet season may be rainfall is vital in moving the applied fertilizer into the soil (Ding et al.,
explained by the heat stress caused by temperatures close to and above 2017). With K, lower uptake was observed during the wet season,
the optimum temperature of 29-30°C (Schlenker & Robers, 2009; considering that this nutrient has a high potential of being lost to the
Schauberger et al., 2017). The studies of Brown and Rosenberg (1999), environment through leaching and runoff, leading to decreased uptake by
Challinor et al. (2014), Bassu et al. (2014), Hussain et al. (2019), and the corn plant (Goulding et al., 2021; Flores et al., 2022).
Anderson et al. (2020) further support the results of the study, as
increases in temperature were noted to negatively impact biomass 4.3 Nutrient Uptake and Biomass Accumulation
accumulation and yield in their paper. In particular, the establishment of
grain sink was found to be impaired by high temperatures, decreasing the The inconsistency between the kernel nutrient uptake and biomass
biomass partitioned to the ear (Lafitte & Edmeades, 1997; Suwa et al., partitioning means that the nutrients (NPK) were absorbed by the kernel,
2010; Rattalino Edreira & Otegui, 2012; Chukwudi et al., 2021). The but did not translate into accumulated biomass. This may be due to the
resulting higher stem weights in the study are also consistent with the increased nutrient uptake in the corn kernel, resulting in a concentration
study of Lizaso et al. (2018), where stem biomass represented a higher greater than its sufficiency range, which leads to luxury consumption, as
percentage of plant biomass during hotter conditions; which is in this shown in Figure 10.
case, during the wet season. Moreover, this non-accumulation to the kernel despite the high
Contrasting literature, such as the study of Rattalino Edreira and nutrient uptake may be caused by other factors influencing plant biomass
Otegui (2013) was also found; where results suggest that although heat such as soil humidity, soil and air temperature, photoperiod, solar
stress severely affects crop production, less effects were particular on the radiation, etc. (Chatzistathis & Therios, 2012; Irving, 2015).
partitioning of biomass to the ear. However, the number of studies
confirming the lower THSC and kernel mean weights during higher
temperatures is found to be more substantial; but this is worth looking
more into, in future studies. In line with this, literature on effects of
increased temperature on the ratio of above- and below-ground biomass
(such as those of Leport et al., 2006; Eziz et al., 2017), were also noted to
be good topics for future research to include the translocation of
assimilates to roots as an effect of high temperature and low rainfall
conditions. In the present study, drought and its interactive effects with
heat stress were not considered, unlike with the research of Lesk et al.
(2016) and Zandalinas (2018), since the daily and cumulative rainfall
depth during the wet season are larger than the values obtained during
the dry season.
The optimum range of daily water requirement of corn at early plant
growth (VE-V5), rapid plant growth (V6-VT), reproductive (R1-R4), and
maturity (R5-R6) stages are 1.91-2.54 mm, 4.57-4.74 mm, 6.1-8.13 mm,
and 2.54-6.10 mm, respectively (Kranz et al., 2008; Alcaide et al. 2019).
Excessive rainfall was observed in the wet season in each of these growth
stages, except for the reproductive stage, where rainfall was insufficient. Fig. 10. Relationship between nutrient concentration in the tissue and crop
In contrast, dry season rainfall was sufficient during the vegetative stage, growth (Havlin et al., 2004).
but insufficient during the reproductive and maturity stages. In terms of
the cumulative rainfall, the optimum range is 390-600 mm (Cakir, 2004; N uptake in the study was the highest in the grain at harvest, and K
Bhandari, 2012; Neild & Newman, 2016). Close to this is the average uptake was highest in the stover at harvest. Hence, a larger impact of the N
cumulative rainfall across sites during the dry season, which was 697 uptake may be observed in the kernel biomass and K uptake in the stover
mm. On the other hand, way above this value was the 939.3 mm rainfall biomass, and the limited impact of the N uptake in the stover biomass and
depth during the wet season. K uptake in the kernel biomass.
Heavy precipitation, as observed during the wet season, has a
significant adverse effect on the biomass accumulation of corn 4.4 Recovery Efficiency in High Temperatures and Heavy Precipitation
(Revadekar & Preethi, 2011; Van der Velde, 2011; Kanchebe Derbile &
Abudu Kasei, 2012; Thornton, 2014; Anderson et al., 2020; Sun et al., Declining grain yields and nutrient use efficiency of corn are among the
2020), whether it may have weaker (Vogel et al., 2019) or stronger impending impacts of climate change, as affected by increases in
association (IPCC, 2007) with yield as compared to temperature. temperature and precipitation. Higher stover and kernel recovery
Partitioning of assimilates to corn ears was affected by high precipitation efficiencies were observed in the study at high temperatures and heavy
events which caused waterlogging, especially during the V3 or third leaf precipitation during the wet season. This phenomenon is inconsistent with
vegetative growth stage of corn (Ren et al., 2014; Hu et al., 2020). the studies of Liang et al. (2013) and Amouzou et al. (2019), predicting a
Consistent with these findings, the results of this study showed decrease in the nutrient use efficiency of crops as affected by future
decreasing biomass in the ear and increasing proportions on the stems climatic conditions, which are described as high in temperature and heavy
and leaves during wet season cropping, as an effect of heavy cumulative in precipitation (IPCC, 2022).
precipitations and excessive rainfall during the critical V3 growth stage of Similarly, the meta-analysis by Yu et al. (2022) found that an increase in
corn. temperature reduced the N recovery efficiency of corn due to the loss of N
in the soil. However, the same study noted that the increase of N recovery
4.2. Soil Water-Nutrient Dynamics and Nutrient Uptake efficiency could also be explained by increasing soil organic content, which
may suggest that this factor, which on average is higher during the wet
Soil nutrient availability is one of the most important factors affecting season for this conducted research, could have affected the recovery
the biomass of crops (Chatzistathis & Therios, 2013). Maximizing the efficiency more significantly than the climatic conditions. Limited studies
10
 K.T. Colarina and P.C. Sta. Cruz
have been conducted on the P and K recovery efficiencies under different were not translated into accumulated biomass, as reflected in the
seasons/climatic conditions, which is an opportunity for future research internal utilization efficiency. This may signify that nutrient uptake
initiatives. exceeded the sufficiency range, inducing luxury consumption. Studies
on the amount of nutrient uptake required to achieve maximum grain
4.5 Kernel Number and Kernel Weight yield with the highest nutrient use efficiency, without reaching luxury
levels, “breaking point” or the toxicity range are recommended. Such
Comparing the results of the yield analysis with the kernel biomass would be beneficial to sustainably increase grain yield to its maximum,
partitioning, it is notable that kernel biomass decreased during the wet without risking environmental degradation due to the over-application
season, while the grain yield increased. This is because grain yield can be of fertilizers.
divided into two components: kernel number and kernel weight. Hence,
despite the decrease in the kernel weight (kernel biomass) during the wet Declaration of Competing Interest
season due to various reasons (e.g. high precipitation during grain filling),
it is entirely possible for the kernel number to increase, resulting to an Kate Mariel T. Colarina has written this paper as part of her MS
ultimate increase in the grain yield (Chen et al., 2017). candidature. She and her thesis adviser, Dr. Pompe C. Sta. Cruz declare that
there are no financial/personal interests or beliefs that affect the
5. Conclusion objectivity and validity of this paper.

The study investigated the effects of seasonal variations on the Acknowledgements

biomass partitioning and nutrient use efficiency of corn, through the
study of the most critical environmental factors to corn production: This thesis would not have been completed without the support and
temperature and rainfall. Based on the comparative analyses of the pre- assistance of several individuals and institutions. Hence, I want to take this
identified yield attributes, biomass partitioning, nutrient uptake, and opportunity to express my utmost gratitude to the following: Dr. Pompe C.
nutrient use efficiency were affected by seasonal variations. The results Sta. Cruz, Dr. Lilia M. Fernando, and Dr. Pearl B. Sanchez, for committing
indicated that during the wet season, which was characterized by higher their valuable time and effort leading to the completion of this thesis;
temperature and precipitation in contrast with the dry season, higher DOST-ASTHRDP-NSC and Project SARAi, as the institutions that provided
nutrient uptakes and nutrient use efficiencies resulted in higher grain academic and financial support to complete this study; and to my loved
yields. Conversely, an inconsistent decrease in kernel biomass ones for their undying love and support. We also thank the editor and
partitioning may have been compensated by higher kernel numbers to reviewers for their constructive suggestions and encouraging comments.
still acquire higher corn yields during the wet season.
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