Volume 11, Number 1 (October 2023):5017-5024, doi:10.15243/jdmlm.2023.111.5017 ISSN: 2339-076X (p); 2502-2458 (e), www.jdmlm.ub.ac.id
Open Access 5017 Research Article
Drought-tolerant lines of Physalis angulata L. improved growth, yield and water use efficiency in drylands
Wiwin Sumiya Dwi Yamika*, Nevy Kusuma Dewi, Budi Waluyo, Nurul Aini, Husni Thamrin Sebayang
Department of Agronomy, Faculty of Agriculture, Brawijaya University, Jl. Veteran, Malang 65145, Indonesia
*corresponding author: [email protected]
Abstract Article history:
Received 3 May 2023 Revised 6 June 2023 Accepted 24 June 2023
Cutleaf groundcherry (Physalis angulata L.) has the potential to be developed in various areas, including dryland. Information on drought-tolerant varieties, lines or genotypes is needed for the development of cutleaf groundcherry in dryland.
Selecting drought-tolerant lines is an alternative for alleviating yield loss potency caused by water shortages. A pot experiment that aimed to investigate the response of cutleaf groundcherry lines to a different level of water deficit, expressed in field capacity (FC), was run in two factors of factorial randomized block design. Each line (PA-01, PA-03, PA-05, PA-08) was set up in water deficit treatment (100, 80, 60, 40, and 20% FC). The result showed that vegetative growth and fruit production, such as fruit number and weight, mainly decreased at 60 or 40 % FC. In contrast, TSS increased at a higher water deficit which was in line with total flavonoid content, even inconsistently. PA-03 and PA-08 experienced a reduction in fruit weight at 40% FC, whereas other lines occurred at 60% FC. Water use efficiency (WUE) increased under severe water stress.
Compared to other lines, PA-03 and PA-08 exhibit higher WUE at 60% FC. In conclusion, PA-03 and PA-08 lines were tolerant of water deficit.
Keywords:
dryland
environmental change lines tolerant
water deficit water use efficiency
To cite this article: Yamika, W.S.D., Dewi, N.K., Waluyo, B., Aini, N. and Sebayang, H.T. 2023. Drought-tolerant lines of Physalis angulata L. improved growth, yield and water use efficiency in drylands. Journal of Degraded and Mining Lands Management 11(1):5017-5024, doi:10.15243/jdmlm.2023.111.5017.
Introduction
Dryland is widely used as an agricultural land where many plants are cultivated. Dryland is often constrained by the lack of water availability in the soil, which can cause plants to be in drought-stress conditions and will cause changes in plant morphology and physiology (Seleiman et al., 2021). Water shortage in dry land might be caused by environmental or climate change impacts. The increasing soil water shortages severity led to degrading soil quality for growing plants. Plants grown under the lack of water are generally smaller than plants that grow normally.
Water is important in the photosynthesis process and the transportation of assimilates to all parts of the plant (Atkins and Smith, 2007). In addition, water functions as a nutrient solvent in the soil and supplies oxygen to the soil (Włodarczyk et al., 2009). In water-
stress conditions, plants tend to experience inhibition of physiological processes in plants and cause leaf stomata closure. This closure is a form of plant adaptation when water is deprived process (Pirasteh‐
Anosheh et al., 2016). Stomata is a place of water loss and carbon dioxide gas (CO2) absorption in photosynthesis. Lower CO2 concentration in the leaf caused by declining stomatal conductance adversely affected the assimilation rate (Ozaslan et al., 2016). In stressed plants, cell division and enlargement are inhibited due to the lack of assimilates produced, causing less assimilate translocation to plant organs, resulting in plant dry weight (Ohashi et al., 2000; Luo et al., 2020). Lack of water during generative growth causes competition between leaves and seeds in utilizing photosynthate, which will cause relatively less fruit to form, small fruit size, and affect the weight and quality of the fruit produced (Nahar and Ullah,
Open Access 5018 2018). In water stress conditions, the plant will exert
to increase water productivity to minimize the hazardous effect on yield. The certain condition of less water supply will increase water use efficiency (Liu et al., 2019). Drought or water shortages in dryland should be restored through sustainable land management and appropriate cultivation. The development of drought-tolerant crops can be an alternative option.
Cutleaf groundcherry (Physalis angulata L.) is a potential crop to be developed in various areas, including dryland. Drought-tolerant lines or varieties are required to improve productivity in dryland.
Information on drought-tolerant varieties or lines is needed to develop cutleaf groundcherry in dryland.
However, that information is still limited. Physalis angulata has become popular nowadays due to its phytochemicals compounds and the beneficial value contained by the plant (Shenstone et al., 2020).
Various parts of the Physalis plant, i.e., fruit, leaf, stem, flower, and root, were reported to have functioned as folk medicine to treat various diseases such as diabetes, stomachache, cancer, malaria, worm, skin disease, infection of fungal, inflammation, etc.
(Kasali et al., 2021).
Cutleaf groundcherry is a potential plant from the Physalis genus; it has a protein content of 27.80%
in fruit and 10.97% in leaves. The fiber content is 10.97% in leaves and 1.83% in fruit. Lipids are 3.67%
in leaves and 2.33% in fruit. While 66.36%
carbohydrates in leaves and 59.70% in fruit (Aliero and Usman, 2016). In addition, other compounds found in cutleaf groundcherry include alkaloids, flavonoids, polyphenols, saponins, physalin, and citric acid (Ayodhyareddy and Rupa, 2016). Various bioactive compounds in the cutleaf groundcherry make this plant widely used as herbal plants (medicines) or consumed directly (fruits).
This research aimed to investigate the response of four lines of cutleaf groundcherry to several levels of water deficit expressed by field capacity.
Materials and Methods Experimental design
The research was conducted from February to May 2020 in a greenhouse located on Jl. Raya Karangan, Donowarih Village, Karangploso District, Malang Regency, East Java. The location is located at an altitude of 550 m above sea level. The air temperature ranges from 26 oC-33 oC with humidity ranging from 60%-75%. This research used a factorial randomized block design consisting of two factors. The first factor was four lines of Physalis angulata L., which were PA 01, PA 03, PA 05, and PA 08. The second factor was the water deficit with a field capacity consisting of 5 levels, i.e., 100, 80, 60, 40 and 20% field capacity.
There were 20 treatment combinations with three replications for each treatment combination.
Plant cultivation
The growing media preparation was done by mixing the soil with cow manure at a dose of 30 t ha-1. The soil was collected from the land near the experiment site.
The mixture media was transferred to a 35 cm x 40 cm polybag weighing 10 kg of each polybag. Planting was done by transferring the seedlings at 21 days after sowing (DAS). The fertilizers used included nitrogen, phosphorous and potassium. They were applied in the dose of 138 kg N ha-1, 36 kg P2O5 ha-1 and 36 kg K2O ha-1 (equal to 300-100-60 kg ha-1 Urea-SP36-KCl).
Urea was applied to equal-split at 7 DAS and 21 DAS, whereas SP36 and KCl were applied fully dose at 7 DAS. Cutleaf groundcherry plants were irrigated daily with a volume of water based on field capacity (FC) treatment. Soil samples were brought to the laboratory to determine soil water content at pF 4.2 and pF 2.5. The volume of water given was measured through the following equation
T = WC . − WC . x Ws Note:
TSWC : Total Soil Water Content WCpF 2.5 : Water content at pF 2.5 WCpF 4.2 : Water content at pF 4.2 Ws : Soil weight
Soil weight at 100% FC was calculated and used to determine other levels of FC by multiplying. The soil weight of each FC level was maintained by rewatering the soil as much as the water evaporated until reaching the weight of their FC condition. Therefore, the soil was weighed before the watering scheduled to estimate the appropriate water irrigated. The volume of water at each watering time was recorded to estimate the water consumption of the overall growth. Pests and diseases were controlled mechanically and chemically by spraying a fungicide (Propineb 70%) and an insecticide (active ingredient Abamectin 18 g L-1).
Plants were harvested on the individual fruit when the calyx turned yellow at once-a-week intervals.
Plant observation
Growth variables were measured at eight weeks after transplanting (WAT). The variables included the number of leaves, leaf area, specific leaf area, and plant dry weight (DW). Several leaves were counted on fully opened leaves of the plant samples. Leaf area measurements were carried out using the ALA method. This method was done by taking leaf samples and calculating the number and the leaf area using LAM (a), then finding the average leaf area of an individual leaf. The formula below can calculate the Leaf area per plant (Widaryanto et al., 2019). On the other hand, SLA and plant DW were measured destructively. They were initiated by taking off the plant sample. Plants were cleaned from the soil attached to the root and oven-dried on each part, i.e., leaf, stem, and root, separately for 48 hours.
Open Access 5019 Plant DW was determined by calculating the total dry
weight of each part (leaf weight (LW)+stem weight (SW)+root weight (RW)). Whereas SLA was calculated by the formula below (Abdelaziz, 2014):
LA = ALA x NL SLA = LA
LW where:
LA : Leaf Area (cm2 plant-1) ALA : Average Leaf Area (cm2 leaf-1) NL : Number of Leaves
SLA : Specific Leaf area (cm2 g-1) LW : Leaf Weight (g)
For yield observation, individual fruit which reached the maturity stage (yellowish to brown of fruit calyx) were picked periodically. Fresh weight was measured on the fruit after taking off the calyx. In the end, the cumulative number and fresh weight of the fruits from the first until the last harvest was served as the number of fruit and fruit weight. Fruit size was evaluated by fruit diameter measurement using a digital caliper.
Fruit properties as a total soluble solid (TSS) were evaluated by dripping fruit juice on a hand refractometer. TSS was expressed in 0Brix. The total flavonoid content of the fruit was analyzed in the laboratory based on the quercetin equivalent method.
Water use efficiency (WUE) of each treatment was evaluated as the ratio between fruit yield (g plant-1) and total water consumed (L plant-1) of overall growth (Chen et al., 2015).
Data analysis
Data recorded from the observations were analyzed using analysis of variance (F test) at the 5% level; if there was a significant effect between treatments, a further test was carried out using the Least Significant Difference (LSD) at the 5% level.
Results and Discussion Soil properties
The soil texture was silt clay loam with a bulk density of 0.86 g cm-3 and a field capacity of 200 mL kg-1. The soil was good enough to hold water and possibly the plant to grow under severe water deficit (20% FC) (Table 1) Fruit yield was also obstructed (Figure 1).
The fertility of the soil was low-medium due to organic C of 0.86%, pH of 5.4, total N of 0.12%, exchangeable K of 0.41 me 100 g-1, and available P of 33 ppm.
Therefore, chemical fertilizers were added in the dose of 138 kg N ha-1, 36 kg P2O5 ha-1 and 36 kg K2O ha-1 Plant growth
Growth variables of cutleaf groundcherry, i.e., number of leaves, leaf area, SLA, and plant DW, were affected by the interaction between lines and water deficit (Table 1). The PA01 and PA03 lines with 80% water supply, the leaf number response was not significantly different from the 100% FC. While the PA05 and PA08 lines with a water supply of 80%-20% FC showed that the number of leaves was not significantly different from that of 100% FC.
Table 1. Growth of cutleaf groundcherry affected by the interaction between lines and water deficit.
Lines Water Deficit (% FC)
Number of leaves
Leaf area (cm2plant-1)
Plant dry weight (g plant-1)
Specific Leaf Area (cm2 g-1)
PA 01 100 93.83 g 3144.41 cde 39.66 h 277.34 cde
80 94.83 g 3527.28 e 40.14 h 274.47 cd
60 54.50 abc 3139.97 cde 25.85 cdefg 387.61 gh
40 52.17 ab 2519.04 bc 20.00 abc 221.40 abc
20 41.67 a 1651.32 a 17.07 a 183.67 ab
PA 03 100 85.50 fg 3195.85 cde 28.62 efg 273.32 cd
80 87.67 fg 3616.82 e 30.45 fg 401.83 h
60 63.33 bcd 2634.50 bcd 26.41 cdefg 310.09 deg
40 58.17 abcd 1892.83 ab 19.03 ab 360.92 efgh
20 57.67 abcd 1400.85 a 16.88 a 263.71 bcd
PA 05 100 75.17 def 3315.35 de 54.69 i 339.71 defgh
80 72.00 cdef 3278.14 cde 49.24 i 265.62 bcd
60 71.00 cdef 3079.95 cde 23.33 abcde 325.16 defgh
40 61.33 bcd 2902.76 cde 22.45 abcde 309.37 defg
20 58.17 abcd 2742.47 cd 21.05 abcd 165.64 a
PA 08 100 84.50 efg 3626.90 e 40.61 h 333.66 defgh
80 84.00 efg 3037.42 cde 32.08 g 378.49 fgh
60 83.33 efg 1700.27 a 31.58 fg 261.97 bcd
40 67.00 bcd 1691.42 a 26.74 deg 297.27 cdef
20 67.00 bcd 1493.72 a 25.16 bcdef 148.75 a
LSD 5% 17.74 761.23 6.69 84.02
Note: Means with different letters indicated significant differences based on LSD 5%.
Open Access 5020 The PA01 and PA03 lines with 80% FC did not show
a decrease in the number of leaves. The number of leaves decreased when the water was given 60%-20%
field capacity. Meanwhile, the PA05 and PA08 lines exhibited a decrease in the number of leaves when water was given 80%-20% FC. Drought stress in plants causes a decrease in the number of leaves which possibly design to reduce transpiration in plants. This might be an adaptive mechanism of the plant toward insufficient water supply (Delfin et al., 2021). In a similar vein, Travlos (2012) reported that water stress in cutleaf groundcherry (Physalis angulata L.) would impact the low level of stem elongation; while water availability is met, the plant will produce more leaves because it has more branches. For plants that are fulfilled their water needs, nutrients are also available so that photosynthesis takes place well and much photosynthate is produced, which is then used for leaf formation.
PA-01 line with 80% FC showed a leaf area response that was not significantly different from 100% FC, the PA-03 line with 80% and 60% FC showed a leaf area response that was not significantly different from 100% field capacity, while the PA05 line with 80%-20% FC showed a leaf area response that was not significantly different from 100% FC;
meanwhile, the PA08 line with 80% FC showed a leaf area response that was not significantly different from giving water at 100% FC. There was no decrease in leaf area in PA-01 and PA-03 lines with a water supply of 80% FC. A decrease in leaf area occurred at a water supply of 60%-20% FC. While PA-05 and PA-08 lines revealed a decrease in leaf area when water irrigated was 80%-20% FC.
Like the plant height, water is needed for cell division and enlargement during the vegetative phase.
Water plays a pivotal role in increasing leaf growth.
The sensitivity of leaves organ toward water deficit was indicated by a reduction in number, length, and width; leaf morphological and physiological characteristics were also reported on Tomatoes (Medyouni et al., 2021) and Cotton (Ödemiş and Candemir, 2022). It is in line with Alves and Setter (2004). They suggested that under water deficit, the leaf produced fewer and smaller cells which disrupted the expansion of the leaf. Leaf expansion was also limited due to reduced turgor pressure (Farooq et al., 2009). Plant DW reduction varied among the lines used. On PA01 and PA05 lines, the reduction started at 60% FC, whereas 80% FC was not different from 100% FC.
Different responses at the lower water content occurred in PA-03 line, where 40% FC started to reduce plant DW. Otherwise, in PA-08 lines, the lightest water deficit had started to reduce plant DW.
The result showed that drought stress constrained assimilate production in photosynthesis due to the limited material used. PA-05 resulted in the highest plant DW at normal (100% FC) and low water deficit (80% FC). It assumed that PA-05 was sensitive to
water deficit (drought) stress. The availability of sufficient water in plants will increase the dry weight of the plant, and conversely, insufficient water will cause a decrease in plant dry weight. Photosynthesis is a physiological reaction constrained by water stress due to some limitations in biochemical factors (Galmés et al., 2007). This led to a tremendous significant reduction in dry matter accumulation (biomass). This result is in congruence with Allahverdiyev and Huseynova (2017). They corroborated that the lack of water reduces plant dry matter due to decreasing photoassimilates and acceleration in plant senescence. Lower plant dry weight might be associated with increasing respiration rate and obstructing biomass accumulation (Akhkha et al., 2011; Khalil et al., 2020).
PA-01 line exhibited an increase in the SLA value at 60 % FC but decreased at 40% FC and lower.
PA-03 response is slightly different, where enhancement occurred at 80% FC and declined at 20%
FC. PA-05 and PA-08 showed a declining SLA value at 20% FC. SLA is the ratio of leaf area to leaf dry weight, which measures the thickness of leaves (Sakya et al., 2015). Lower SLA described the reduction in dry matter production per unit leaf area and correlated negatively with leaf thickness (Hummel et al., 2010;
Abdelaziz, 2014).
Yield and quality attributes
A number of fruits were not affected by the interaction between lines and water deficit. A number of fruits were mainly influenced by watering treatment (Table 2). Under 80 and 60% FC, the number of fruits was not different from 100% FC. The reduction of fruits number occurred at 40 and 20% FC. Provision of water at 100% FC shows a high number of fruit compared to other field capacity water supply. The number of fruit decreases along with the smaller field capacity. This suggests that plants with water shortages tend to produce less and less fruit, also the other way around.
The reproductive stage of fruit crop starts from flowering and need sufficient supply (Fischer et al., 2016). Fruit number was a yield component that was reduced under a high level of water stress (Alomari- Mheidat et al., 2023). Fruit weight was affected by the interaction between the use of lines and water deficit (Figure 1a.). PA-01 and PA-05 resulted in lower fruit weights starting at 60% FC, PA03, and PA08 started at 40% FC. Under 80 and 60% FC, those PA-03 and PA- 08 were able to produce fruit as good as 100% FC. If observed on each water deficit level, under 100% FC, PA-05 resulted in higher fruit weight over PA-01 and PA-03 but was not significantly different from PA-08.
It tends to be similar to the 80% FC condition, where PA-05 was still higher than PA-03 but was not different from PA-01 and PA-08. On the contrary, when plants were exposed to water deficit at 60% FC, PA-03 and PA-08 resulted in higher fruit weight compared to PA-05. In the higher water deficit level, i.e., 40 and 20% FC, lines did not affect fruit weight.
Open Access 5021 Table 2. Fruit production and fruit properties of cutleaf groundcherry lines and the effect of water deficit.
Treatments Number of Fruits (plant-1) Fruit Diameter (mm) Total Soluble Solid (0Brix) Lines
PA-01 25.53 14.18 10.28
PA-03 22.40 14.60 9.44
PA-05 23.93 14.38 9.79
PA-08 26.13 14.80 9.63
LSD 5% ns ns ns
Water Deficit (% FC)
100 29.25 b 15.29 c 9.13 a
80 27.91 b 15.26 c 9.66 ab
60 23.41 ab 14.54 bc 9.86 ab
40 21.25 a 13.95 ab 10.11 b
20 20.66 a 13.39 a 10.15 b
LSD 5% 6.52 1.07 0.95
Note: Means with different letters indicated significant differences based on LSD 5%, ns = not significant.
Figure 1. Fruit weight (a) and total flavonoid (b) of cutleaf groundcherry fruit affected by the interaction between lines and water deficit. Means followed by different letters on each bar indicate significant differences
based on LSD 5%.
fgh
cdef
h fgh gh
def
gh
cde def efg
abc
efg bcde
ab ab abc
abcd
ab a a
0 20 40 60 80 100
PA-01 PA-03 PA-05 PA-08
Fruit Weight (g plant-1)
Treatments
100 80 60 40 20
Water deficit (% FC) a
gh
d d
bc e
c
h d d
ab
k
h d
a
ef
i
fg ef
d
j
0.5 1.0 1.5 2.0 2.5 3.0 3.5
PA-01 PA-03 PA-05 PA-08
Total Flavonoid (mg g-1)
Treatments
100 80 60 40 20
Water deficit (% FC) b
Open Access 5022 Figure 2. Water Use Efficiency (WUE) of cutleaf groundcherry growth affected by the interaction between lines
and water deficit. Means followed by different letters on each bar indicate significant differences based on LSD 5%.
This confirmed that a water deficit at 40% was a critical level for cutleaf groundcherry, where a reduction in yield occurred in all lines. Water stress during generative growth decreased fruit weight, which might be associated with insufficient carbon demand and limited water resources for photosynthesis, which caused lower carbohydrate results (Berman and Dejong, 1996). This condition led to the competition for limited carbon sources, which caused relatively less fruit production in its number, size, and weight.
Fruit properties, fruit diameter, and TSS were not affected by the interaction between the use of lines and water deficit with field capacity. Regardless of the line, the provision of water with field capacity significantly affected fruit diameter and TSS (Table 2). Fruit diameter was reduced by under 40 and 20% FC. Fruit diameter is getting smaller along with the provision of water with a smaller field capacity. Plants with adequate water requirements tend to produce larger fruit diameters. The final size of the fruit was affected by cell division in the stage of fruit development, and water deficit negatively affected fruit size (Bertin, 2005). In addition, fruit development as a sink organ was affected by carbohydrates which might be controlled by the interaction between the line and the environment (Prudent et al., 2010). On the other hand, the water deficit had a positive effect on TSS. Under 20 and 40% FC, TSS was higher compared to 100%
FC. It means that low water content in the soil produced high TSS, which can be associated with fruit sweetness (Sucrose). Increasing TSS under water stress conditions was also reported on Tomato fruit (Alordzinu et al., 2022). Soluble sugar was one of the osmotic compounds. The accumulation positively correlated to water deficit. Water deficit-affected genes involved in fruit quality underlined the mechanism of increasing soluble solid accumulation in
the fruit (Bai et al., 2023). The total flavonoid content of cutleaf groundcherry fruit showed a wide variation among lines and water deficit treatment and was affected by their interaction (Figure 1b). Generally, the highest total flavonoid content was obtained from PA- 05, which was treated with 60% FC. Whereas PA-08 produced higher total flavonoid content in line with increasing water deficit (lower FC). On each line, total flavonoid content tended to increase in certain water deficit levels and exhibit a decline in other water deficit levels. Similarly, it was reported by Jin et al.
(2022) in tomato fruit where flavonoid content was found to be higher under the substrate's 80-65%
moisture capacity.
Water use efficiency (WUE)
The interaction between line and water deficit treatment influenced WUE significantly (Figure 2.).
PA-01 with 80% and 60% FC showed a response similar to giving 100% field capacity water, and PA03 and PA08 lines with 80% FC water distribution showed a response similar to giving 100% field capacity water. The PA05 line with 80%-40% water supply showed a significantly different response to 100% water supply field capacity. All lines PA01, PA03, PA05, and PA08 showed the highest water use efficiency value at 20% FC. The efficiency of water use is the ratio between plant yield (fruit weight) and water requirements and shows the ability of plants to convert water into yield (Chen et al., 2015). The results showed that providing water with a smaller field capacity could increase water use efficiency. Ismail (2010) reported that WUE tends to be increased under water deficit in bird pepper plants. Higher WUE at the same soil water content implied more yield obtained per water unit and was attributed to drought tolerance.
However, increasing WUE for water resource saving should not sacrifice a tremendous yield of lost potency.
abc a
abcdefg
ab
abcdef abcde
hi
abcd
abcdef fgh
bcdefgh cdefg
ij
defg
ghi efgh
l
k
l
j
0 4 8 12 16 20
PA-01 PA-03 PA-05 PA-08
WUE (g fruit FW L-1)
Treatments
100 80 60 40 20
Water deficit (%FC)
Open Access 5023 Conclusion
A high water deficit (≤40% FC) significantly reduced the growth and yield of all cutleaf groundcherry lines used. Generally, cutleaf groundcherry has a good adaptation under a low water deficit and varied response under an increasing water deficit level. PA- 05 have better growth and yield when planted under normal condition. However, it turned out to be lower when planted at a high level of water deficit (≤60%
FC). PA-03 and PA-08 have a potency for growing on dryland which experienced drought. It was evidenced by plants' ability to produce better biomass and fruit yield under a moderate water deficit (60% FC). It can be attributed to the higher WUE at the level of water deficit. For fruit quality consideration, PA-08 should be advanced due to the high total flavonoid content obtained at a high level of water deficit.
Acknowledgements
This study was supported by the Ministry of Research, Technology, and Higher Education through the Competitive
Research Grant by contract number
292.13/UN10.C10/PN/2020 and the Indonesian Ministry of Education, Culture, Research, and Technology, through BPPDN scholarship.
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