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Effect of waterlogging on growth and development of selected green gram varieties   

 

 

 Samson Onyango 
Ochar1+ 
Phoebe Anyango 
Sikuku2 
John Collins Onyango3 

1,2,3Department of Botany, Faculty of Biological and Physical Sciences, Maseno 
University, Kisumu-3275-40100, Kenya.  
1Email: ochars2008@gmail.com  
2Email: sphoebe@maseno.ac.ke  
3Email: jconyango@yahoo.com  

 

 
(+ Corresponding author) 

 ABSTRACT 
 
Article History 
Received: 18 April 2025 
Revised: 4 July 2025 
Accepted: 14 July 2025 
Published: 21 July 2025 
 

Keywords 
Chlorophyll content  
Green grams 
Physiological responses  
Vegetative stage 
Waterlogging 
Yield components.  

 
 

 
This experiment studied the effect of waterlogging on the growth and development of 
selected green gram varieties to recommend the variety that can withstand waterlogging. 
The seeds of three selected green gram varieties were planted in 10-litre capacity pots 
arranged in a completely randomized design. Waterlogging was induced at the 
vegetative stage, 21 days after sowing (DAS), by dipping 10-litre pots into larger 20-litre 
pots and maintaining standing water at 3cm above the soil level for 3 days (T3), 6 days 
(T6), and 9 days (T9). The control (T0) was achieved by watering the pots with 500ml 
of water per day. Net assimilation rate, transpiration rate, and stomatal conductance were 
measured using LI-COR 680 portable photosynthesis systems. Chlorophyll content was 
determined using the Arnon method. Measurements were taken from three plant samples 
per variety regularly throughout the study period. KAT 00301 and KAT 00309 
maintained significantly higher net assimilation rates, stomatal conductance, and 
chlorophyll content than KAT 00308 under waterlogging treatments. KAT 00308 was 
more vulnerable to waterlogging; the variety wilted and died under prolonged 
waterlogging, but KAT 00301 and KAT 00309 survived, though with reduced yield. 
Both small and commercial green gram farmers should plant KAT 00301 and KAT 00309 
to minimize losses due to waterlogging.  
 

Contribution/Originality: Green gram, being a drought-tolerant crop, has limited information regarding the 

effects of waterlogging on its performance. The study on the impact of waterlogging on the growth and development 

of newly released varieties by KALRO aims to identify varieties that can withstand waterlogging conditions, thereby 

giving the study an originality status. 

 

1. INTRODUCTION 

Green gram is the third most important pulse crop after chickpea and pigeon pea. Ecologically, green grams 

require an altitude of 0-1600m above sea level, sandy loam and clay soils with a pH range of 5.5-7.5. It is tolerant to 

drought and has an annual rainfall requirement range between 350-700mm. It has a well-developed root system with 

tap roots and lateral roots for water absorption when limited. Being a reliable protein source, green gram is a stable 

food security source. Mature seeds of green gram contain proteins, carbohydrates, minerals, fibers, as well as 

antioxidants [1]. Globally, in countries where meat is culturally prohibited from being used as food, such as India, 

green grams offer a reliable source of protein [2]. Apart from offering a stable source of food and nutritional security, 

green gram is an important income source for many rural households. Kanavi et al. [3] reported that despite the 

economic importance of green grams, abiotic and biotic stresses have caused an inordinate decline in its overall 

Current Research in Agricultural Sciences 
2025 Vol. 12, No. 2, pp. 115-122 
ISSN(e): 2312-6418 
ISSN(p): 2313-3716 
DOI: 10.18488/cras.v12i2.4307 
© 2025 Conscientia Beam. All Rights Reserved. 

 
 
 

 
 
 
 

 

 
 
 
 

mailto:ochars2008@gmail.com
mailto:sphoebe@maseno.ac.ke
mailto:jconyango@yahoo.com
https://orcid.org/0009-0006-2304-8797
https://www.doi.org/10.18488/cras.v12i2.4307


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116 

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production. Green gram has the potential to produce 2.5–3.0 tons per hectare. Despite this potential, due to abiotic 

and biotic constraints, the productivity stagnates at 0.5 tons/hectare [4]. Green gram thrives well in semi-arid land 

in Kenya, where it is cultivated for both subsistence and commercial purposes. In Machakos, Kitui, Tharaka-Nithi, 

and Makueni counties, green gram production has proven to be more successful, with Machakos County leading in 

green gram production [5].  The newly released varieties of green gram by the Kenya Agricultural and Livestock 

Research Organization (KALRO) include KAT 00301, KAT 00308, and KAT 00309. They are commonly cultivated 

in most parts of Kenya. All the varieties have a short maturity period and perform well in the arid and semi-arid lands 

of Kenya. Table 1 shows the physical characteristics of the selected green gram varieties.  

 

Table 1. Physical characteristics of  the newly released green gram varieties. 

Green gram variety  Physical properties Days to maturity Yield in bags /ha 

 KAT 00301 • Grains are green and shiny in color  

• Grain size (6-7g/100 seeds) 

60 - 70 1800-2300kg/ha 

KAT 00308 • Grains are shiny green in color 

• Grain size (8-10g/100 seeds) 

65-75 1800-2100 kg/ha 

KAT 00309 • Grains are green and shiny in color  

• Grain size (8-10g/100 seeds) 

60-65 1800-2100 kg/ha 

 

The early maturity enables them to survive terminal droughts in arid and semi-arid areas. Since green gram is a 

drought-tolerant crop, its performance in waterlogged soils remains questionable. One of the prevalent problems 

during the growing period of green grams is waterlogging [6]. The physiological and chlorophyll content responses 

of the aforementioned green gram varieties to waterlogging are yet to be fully understood. Studies on the effect of 

waterlogging on green grams have been explored, and most of these studies focused on yield components and a single 

growth parameter [5, 7-9]. According to studies by Ahmed et al. [9], in particular, it was shown that green grams 

have the ability to recover from short-term waterlogging damage, and that the response to waterlogging depends on 

the variety. Amin et al. [8] reported that the survival rate of green gram genotypes was less than 20%. Kumar et al. 

[10] reported that green gram varieties subjected to waterlogging for nine days lost their photosynthetic apparatus 

by more than 80% and did not recover from waterlogging damage. The above conclusions were arrived at by 

analyzing at least one physiological parameter. Since physiological parameters and chlorophyll development directly 

influence morphology and yield, it is important to investigate further how waterlogging affects the physiological 

parameters and chlorophyll content of the newly released varieties of green grams, particularly in Kenya, where such 

studies have remained relatively scanty. 

More than 90% of agricultural systems in Kenya are rain-fed; thus, they are highly vulnerable to changes in 

climate. There is an overall shift in rainfall distribution, with floods becoming more likely than the opposite extreme 

[11]. The rainfall patterns in Kenya have been changing, which has been attributed to climate change caused by 

global warming. Fluctuations in rainfall due to unpredictable and erratic patterns expose rain-fed agricultural systems 

in Kenya to increased vulnerabilities. Prolonged flooding impacts the physiological parameters and chlorophyll 

content of green grams. The response of physiological and chlorophyll content to waterlogging manifests in the 

morphology and yield components of green grams. A shift in water regimes during early growth stages destabilizes 

crop development and negatively influences subsequent stages, ultimately affecting final yield. Therefore, it is 

essential to assess the physiological and chlorophyll content responses of selected green gram varieties to 

waterlogging and identify varieties capable of withstanding such conditions. The study was conducted to determine 

the effects of waterlogging on transpiration rate, stomatal conductance, net assimilation rate, and chlorophyll content 

of selected green gram varieties, with the aim of recommending suitable varieties to farmers and plant breeders for 

further breeding efforts towards waterlogging tolerance. 

 



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2. MATERIALS AND METHODS  

2.1. Experimental Site and Climate 

The research was conducted at the University Botanic Garden in Maseno, Kenya, under greenhouse conditions. 

The university is located in the western region of Kenya, along the equator. The climate is hot and humid, with 

temperatures ranging from 660F to 830F. 

 

2.2. Experimental Design and Treatments 

The experiment was conducted using a completely randomized design with four treatments, each having three 

replications. The treatments were as follows: 

T0- control.  

T3- Three days of water logging.  

T6- Six days of water logging.  

T9- Nine days of water logging.  

 

2.3. Experimental Layout  

The research experimental unit consisted of a ten-litre capacity pot. The pots were perforated and filled with 

solarized soil collected from the University Botanic Garden. The soil was collected using a jembe and soil auger and 

was mixed with two teaspoonfuls of DAP fertilizer per pot to hasten root establishment. The seeds of the three 

varieties, KAT 00301, KAT 00308, and KAT 00309, were obtained from KALRO-Kisumu. 

 

2.4. Sowing and Crop Management  

Ten seeds of each variety were planted per pot, and thinning to three was performed fourteen days after sowing. 

From the day of sowing, each pot was watered with 500 ml of water daily. At 21 DAS, waterlogging was induced for 

three successive days for treatment one (T3), six successive days for treatment two (T6), and nine successive days for 

treatment three (T9). The treatments were modified from Amin et al. [12]. The waterlogging status was achieved 

by immersing 10-litre pots containing green gram crops into larger 20-litre pots, and subsequently maintaining 

standing water at 3 cm above the soil surface in each pot for the specified number of days in each treatment. This 

method was modified from  Amin et al. [12] and Ahmed et al. [9]. Each treatment was replicated three times, and 

the pots were arranged in a completely randomized design (CRD). This was in accordance with Amin et al. [12]. 

Control treatment was achieved by watering each pot with 500 ml of water per day throughout the growing period 

of the green grams. After the termination of waterlogging, the 20-litre pots were removed, and each pot was watered 

with 500 ml of water per day throughout the growing period. This was modified from Sosiawan et al. [13]. Weed 

control was achieved by uprooting them from the pots.  

 

2.5. Data Collection 

The data on net assimilation rate, transpiration rate, and stomatal conductance were measured using LI-COR 

680 portable photosynthesis systems, and chlorophyll content was determined using the Arnon method. The data 

were collected from three plant samples of each variety for every treatment throughout the study period after 

initiating the treatments. 

 

2.6. Data Analysis 

The data were subjected to analysis of variance using SAS (version 9.1), and separation was performed using the 

LSD test at a 5% significance level. 

 

 



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3. RESULTS  

3.1. Net Assimilation Rate  

Three days, six days, and nine days of waterlogging significantly reduced the net assimilation rate of all three 

green gram varieties (p≤0.05). At three days of waterlogging, there was a significant difference in the net assimilation 

rate among the three varieties (p≤0.05), with KAT 00309 maintaining a higher rate of net assimilation compared to 

KAT 00301 and KAT 00308. At six days of waterlogging, there was no significant difference in the reduction of the 

net assimilation rate among the three varieties (p≥0.05). At nine days of waterlogging, KAT 00301 maintained a 

significantly higher net assimilation rate compared to KAT 00309 (p≤0.05). Table 2 shows the effect of different 

waterlogging regimes on the net assimilation rate (µmol·m-2s-1) of three green gram varieties. The letters a, b, and c 

indicate significant differences between the varieties for each treatment. Means with the same letter for a particular 

treatment are not statistically significantly different. 

 

Table 2. Effect of  different waterlogging regimes on net assimilation rate (µmol-2s-1) of  the three green gram varieties.  

 T0 T3 T6 T9 

KAT 00309  19.336a 6.1371a 0.5394a 0.5166a 

KAT 00308 8.981b 2.4318b 0.4592a 0.0000c 

KAT 00301 9.484b 0.5039c 0.44084a 1.6048b 

LSD 3.6989 1.3911 0.4445 0.3977 
Note: Means with the same letters along a column are not significantly different (p≥0.05). The values represent the means of three replicates. 

 

3.2. Stomatal Conductance 

Waterlogging significantly reduced stomatal conductance of all the green gram varieties (p≤0.05).  

At three and six days of waterlogging regimes, KAT 00309 and KAT 00301 exhibited higher rates of stomatal 

conductance than KAT 00308; however, the rates of stomatal conductance among the three varieties were not 

significantly different (p≥0.05). At nine days of waterlogging regime, KAT 00301 had a significantly lower rate of 

stomatal conductance than KAT 00309 (p≤0.05). During the six and nine days waterlogging regimes, all plants of 

the KAT 00308 variety had withered. Table 3 presents the effect of waterlogging on stomatal conductance (mol-2s-

1) of the three green gram varieties. The letters a, b, and c indicate significant differences between the means of the 

varieties for each treatment. Means with the same letter for a particular treatment are not statistically significantly 

different. 

 

Table 3. Effect of  different water logging regimes on stomatal conductance(mol-2s-1) of  the three green gram varieties.  

 T0 T3 T6 T9 

KAT 00309  0.69840a 0.05498a 0.05263a 0.06793c 

KAT 00308 0.12557b 0.02186a 0.01476a 0.00000b 

KAT 00301 0.51158a 0.05342a 0.04039a 0.032302a 

LSD 0.206 0.0415 0.0512 0.0229 
Note: Means with the same letters along a column are not significantly different (p≥0.05). The values represent the means of three replicates. 

 

3.3. Transpiration Rate 

Waterlogging significantly reduced the rate of transpiration in all green gram varieties, with nine days having 

the greatest impact (p≤ 0.05). At three days of waterlogging and six days of waterlogging, there was a significant 

difference in the reduction of transpiration rate between KAT 301 and the other two varieties (p≤0.05). At nine days 

of waterlogging, there was no significant difference in the reduction of transpiration rate between KAT 00301 and 

KAT 00309 (p≥0.05). At six days and nine days of waterlogging regimes, all the KAT 00308 plants withered. Table 

4 presents the effect of waterlogging on the transpiration rate (mol m-2s-1) of the three green gram varieties. The 

letters a, b, and c indicate significant differences between the varieties for each treatment. Means with the same letter 

for a particular treatment are not statistically significantly different. 



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Table 4. Effect of  different water logging regimes on transpiration rate(mol-2s-1) of  the three green gram varieties.  

 T0 T3 T6 T9 

KAT 00309  0.0169621a 0.003835b 0.0006570b 0.0005685a 

KAT 00308 0.0070334b 0.001411b 0.0004608b 0.000000b 

KAT 00301 0.0143091c 0.004026a 0.0034330a 0.0006357a 

LSD 0.0018 0.0026 0.0015 0.0005 
Note: Means with the same letters along a column are not significantly different (p≥0.05). The values represent the means of three replicates. 

 

3.4. Chlorophyll Content 

Waterlogging significantly reduced the chlorophyll content in all green gram varieties (p≤0.05). At three days 

of waterlogging, KAT 00309 had a significantly higher chlorophyll content compared to KAT 00308 and KAT 00301 

(p≤0.05). At six days of waterlogging, chlorophyll content among the three green gram varieties was not significantly 

different (p≥0.05); however, KAT 00309 recorded a higher chlorophyll content compared to the other varieties. At 

nine days of waterlogging, KAT 00301 had a significantly higher chlorophyll content than KAT 00309 (p≤0.05). At 

this stage, KAT 00308 had withered and died. Table 5 presents the effect of waterlogging on chlorophyll content 

(mol-2s-1) of the three green gram varieties. The letters a, b, and c indicate significant differences between the 

varieties for each treatment. Means with the same letter for a particular treatment are not statistically significantly 

different. 

 

Table 5. Effect of  different water logging regimes on chlorophyll content of  the three green gram varieties (mg m-2).  

 T0 T3 T6 T9 

KAT 00309 19.336a 6.1371a 0.5394a 0.5166 b 

KAT 00308 8.981b 2.4318 b 0.4592a 0.0000b 

KAT 00301 9.484b 0.5039 c 0.4408a 1.6048a 

LSD 3.6989 1.3911 0.4445 0.6288 
Note: Means with the same letters along a column are not significantly different (p≥0.05). The values represent the means of three replicates. 

 

4. DISCUSSION  

4.1. Net Assimilation Rate 

The rate of net assimilation in all green gram varieties significantly decreased under different waterlogging 

conditions (p≤0.05). This is consistent with previous findings of Prasanna and Ramarao [14] who reported that four- 

and six-day waterlogging significantly lowered net assimilation rate in pigeon pea genotypes. However, the results 

are not consistent with findings of Oo et al. [15] who reported that the net assimilation rate in green gram genotypes 

was less affected by flooding. Reduction in net assimilation rate is attributed to reduced stomatal conductance and 

reduction in leaf area. Reduced stomatal conductance reduces the rate of diffusion of carbon dioxide into the mesophyll 

cells, thus lowering carbon dioxide assimilation. Lowered net assimilation rate could also be linked to chlorosis, which 

leads to loss of photosynthetic pigment, hence reducing the surface area for photosynthesis. An increase in the 

duration of waterlogging increased chlorosis in all green gram varieties. According to Islam et al. [16] deduction in 

photosynthetic activity in green gram genotypes under waterlogging conditions is due to both stomatal and non-

stomatal limitations. 

The study revealed that KAT 00308 was more vulnerable to waterlogging, hence it had the lowest rate of carbon 

dioxide assimilation. KAT 00301 and KAT 00309 maintained slightly higher rates of carbon dioxide assimilation 

under waterlogging stress. KAT 00308 lacks mechanisms to counter chlorophyll degradation, hence it experienced 

rapid chlorophyll degradation that greatly limited photosynthetic activities. Moreover, KAT 00301 and KAT 00309 

developed mechanisms to counter rapid chlorophyll degradation; therefore, photosynthetic activities were not greatly 

affected. 



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4.2. Stomatal Conductance 

Waterlogging significantly reduced stomatal conductance in all three green gram varieties. This confirmed the 

findings of Islam et al. [16] and Takele and McDavid [17]. Reduction in stomatal conductance under waterlogging 

stress is a mechanism employed by plants to curtail transpiration. This is supported by the fact that waterlogging 

stress in all green gram varieties under study significantly lowered the rate of transpiration. Waterlogging may have 

led to the accumulation of ABA, which induced stomatal closure, hence increasing stomatal resistance. High stomatal 

conductance resistance may also be attributed to the lack of oxygen availability in the soil around the root hairs, 

which hampers the normal functioning of roots. 

Among the three green gram varieties, there was a significant difference in stomatal conductance under different 

waterlogging conditions. KAT 00308 had a significantly lower rate of stomatal conductance compared to KAT 00301 

and KAT 00309. This indicates that KAT 00308 is highly vulnerable to waterlogging. Significantly higher stomatal 

conductance in KAT 00310 and KAT 00309 suggests that these two varieties may withstand waterlogging due to 

mechanisms that regulate stomatal conductance. 

 

4.3. Transpiration Rate  

The rate of transpiration significantly decreased in all green gram varieties with an increase in days of 

waterlogging (p≤0.05). The reduction in the rate of transpiration with increased duration of waterlogging is 

associated with increased stomatal closure. This is consistent with the findings of Worku [18], who reported that an 

increase in waterlogging duration significantly decreased the rate of transpiration in green grams. Takele and 

McDavid [17]  also reported that an increase in waterlogging period decreased the rate of transpiration in pigeon 

pea genotypes, while Ahmed et al. [9] reported that 8-day waterlogging sharply declined the rate of transpiration in 

mung bean genotypes. 

There was a significant difference in transpiration rates among the three green gram varieties. KAT 03008 had 

a significantly lower transpiration rate compared to KAT 00301 and KAT 00309 under waterlogging treatments. 

The significantly low rate of transpiration in KAT 00301 under waterlogging is attributed to rapid stomatal closure. 

Similarly, the higher rates of transpiration in KAT 00301 and KAT 00309 under waterlogging could be attributed to 

reduced stomatal closure. 

 

4.4. Chlorophyll Content 

Waterlogging significantly reduced chlorophyll content in all green gram varieties. The reduction was more 

pronounced with an increase in the duration of waterlogging. This is in agreement with the findings of Prasanna and 

Ramarao [14] in green grams and Kumar et al. [10] in green grams; a decrease in chlorophyll content may be 

attributed to chlorophyll degradation and chlorosis. A reduction in chlorophyll content may also be linked to a 

decrease in nitrogen content of leaves due to the cessation of biological nitrogen fixation as a result of waterlogging. 

Among the three green gram varieties studied, KAT 00308 exhibited significantly lower chlorophyll content 

compared to KAT 00301 and KAT 00309 under waterlogging treatments. The loss of chlorophyll was evidenced by 

increased yellowing of leaves, which was eventually followed by wilting and death of KAT 00308 under six- and nine-

day waterlogging regimes. Faster chlorophyll degradation in KAT 00308 may be attributed to the almost immediate 

cessation of nitrogen uptake under waterlogging conditions. Significantly higher chlorophyll content in KAT 00301 

and KAT 00309 under waterlogging conditions suggests that nitrogen uptake in these varieties is not greatly affected 

by waterlogging. 

 

5. CONCLUSION  

Notably, KAT 00308 maintained a relatively low net assimilation rate, low transpiration rate, and low stomatal 

conductance under waterlogging regimes compared to KAT 00301 and KAT 00309. Similarly, KAT 00308 exhibited 



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faster chlorosis, while KAT 00301 and KAT 00309 maintained higher chlorophyll content under waterlogging 

regimes. Six-day and nine-day waterlogging treatments were detrimental to KAT 00308; hence, all plants wilted and 

died, failing to reach the reproductive stage. The variety, therefore, cannot withstand prolonged flooding conditions. 

KAT 0301 and KAT 309 survived six- and nine-day waterlogging, though with reduced final yield. The two 

varieties showed some degree of tolerance to waterlogging, with KAT 00309 less affected physiologically. Both small 

and commercial green gram farmers should plant either KAT 00301 or KAT 00309 to minimize losses due to 

waterlogging. Moreover, either KAT 00301 or KAT 00309 could therefore be better varieties to be improved through 

genetic experiments to develop varieties that can withstand waterlogging for continued green gram production 

throughout all seasons. 

 

Funding:   This study received no specific financial support. 
Institutional Review Board Statement: Not applicable. 
Transparency: The authors state that the manuscript is honest, truthful, and transparent, that no key aspects 
of the investigation have been omitted, and that any differences from the study as planned have been clarified. 
This study followed all writing ethics. 
Competing Interests: The authors declare that they have no competing interests.  
Authors’ Contributions: All authors contributed equally to the conception and design of the study. All 
authors have read and agreed to the published version of the manuscript. 

 

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