Keywords
Drip, rice, phenotyping, root, yield
This article is included in the Agriculture, Food and Nutrition gateway.
Drip, rice, phenotyping, root, yield
Aerobic rice refers to rice grown under non-water stagnation. The growth of the root system in rice is restricted under aerobic conditions, which is the reason for poor yield (Kato et al., 2010). In aerobic soil conditions, the soil’s interaction with rice is primarily focused on the root system. Therefore, the root system is the first barrier to face stress found in aerobic soil conditions. Water management of rice crops is a sensitive tool in aerobic rice cultivation practice, which has been demonstrated by alteration in rice root anatomy (Mostajeran & Rahimi-Eichi, 2008).
A deeper root system of rice eases water stress and improves the uptake of nutrients and water in deep soil layers (Lilley et al., 1994). Rice cultivars with deep rooting and higher root density are much more favorable to aerobic conditions than lowland conditions (Matsuo et al., 2010).
Regulation of root traits to aerobic conditions could be attained by phenotyping for root traits and rhizosphere management by application of water and nutrients to the root zone (Zhang et al., 2010). Drip irrigation stimulates fibrous root production with specific changes in root system architecture (Raj et al., 2013). Similarly, drip irrigation with humigation plants shows favorable root growth with grain yield in rice (Vanitha & Mohandass, 2013).
This study focuses on phenotyping of root traits, such as root length, density, and distribution, under various drip treatments, related to the root response of rice genotypes. Consequently, the phenotyping of root traits and grain yield under drip environment was analyzed by the present study.
A root phenotyping experiment was conducted during the summer season of 2013 and 2014, using JKRH 3333, TNRH 180 and ADT(R)45 as the test rice varieties at Tamil Nadu Agricultural University (Coimbatore, Tamil Nadu, India). Seeds were manually sown in the field at 20×10cm spacing. The open pan evaporation (PE) values (125% pan evaporation) were used to calibrate irrigation scheduling, and drip irrigation was supplied via pipe at 40mm outside diameter (OD) by 7.5HP motor with a pressure of 1.5kg/cm-2 from a bore well. Solar powered and well-operated drip irrigation sources, 0.8 and 1.2m lateral distances, 5–10 and 15–20cm depth sub-surface drip (SDI) were the treatments adopted at field level. The conventional aerobic practice was scheduled at 1.25 irrigation water (IW)/cumulative pan evaporation (CPE) ratio to 3.0cm. It was named as conventional aerobic rice. The recommended dose of 150:50:50 kg:ha-1: NPK water soluble fertilizers were used to fertigate the crops using the Venturi flume at weekly intervals. Further information on genotypes, experimental set up and fertigation schedule are given in Supplementary File 1.
Root length was estimated during the flowering phase (80 days after sowing) from core samples (Kato et al., 2006). Rice roots were removed carefully from the soil by root auger without damaging the roots. After the samples were oven-dried at 80°C for 72h, root lengths and weights were measured. Root length (m hill-1) = sample root length (cm) × total root weight (g)/sample root weight (g). Root dry weight was expressed as g/hill. The specific root length was estimated as a ratio by root length to root dry weight. Four soil cores (50mm diameter, 35cm depth) per plot were taken next to the plant and between the rows (20cm) with a soil sampler. Cores of soil were separated into 0–15 and 15–35cm, then washed using water and sieved by 0.5mm mesh sieve. The root length (RLD), as well as root mass density (RMD), was determined using the formula of Pantuwan et al. (1997), and the values were expressed as cm/cm3 and mg/cm3 of the soil, for RLD and RMD respectively. Root volume was recorded using the water displacement technique (Bridgit & Potty, 2002) and expressed as cm3/hill.
Core sampled roots were washed thoroughly and dehydrated using 80, 90 and 100% alcohol. Dehydrated roots were embedded in slides using paraffin. Slides were kept for imaging the root system using camera (Sony 12.1 megapixel) mounted Leica D1000 microscope at 10X magnification (Guo et al., 2008).
Adenosine triphosphatase (ATPase) activity of the root was assayed according to Wayne (1955) at 32°C using ATP (sodium salt) as a substrate, and the reactions were terminated by the addition of 2.0 mL cold 10% trichloroacetic acid. The ATPase enzyme activity was expressed as µg Pi g-1 h-1.
Harvesting of the crop (grain) was performed from the net plot level (2.4×7.0m) at 120 days after sowing. The yield of rice grain was calculated to hectares at 14% moisture level and expressed as kg ha-1. The filled grain percentage (%) was calculated by the ratio of total filled grain with total spikelet numbers in panicles. The dry weight of grain and total grain dry weight per hill ratio was used to measure the harvest index (HI; %) at harvest stage of the crop (Yoshida et al., 1971)
The recorded mean data were analyzed with AgRes software (version 7.01) ANOVA (Analysis of Variance) package for researchers 1994, Pascal Intl software solutions. Significance was assessed at 95% (p<0.05) and 99% (p<0.01) confidence level (Gomez & Gomez, 1984). F values were calculated using the method as described in http://www.biokin.com/tools/fcrit.html.
Total root length (TRL) is the size of the total root system, which is the major determinant for water and nutrient uptake. The drip irrigation system used in the present study in aerobic rice showed significant variation among root traits. Regarding the TRL for the different rice genotypes, a longer length was observed in with JKRH 3333 (6.2m/hill), followed by TNRH 180 (50.7m/hill) and ADT(R) 45 (38.6m/hill) (Table 1).
Among the genotypes, a significantly higher root volume was observed in JKRH 3333 (66cm3/hill), followed by TNRH 180 (61.7cm3/hill) and ADT(R)45 (53.2cm3/hill). From the main plot treatment, the solar drip irrigation recorded an increased root volume of 43.4% compared with the well-operated drip irrigation treatment.
LD: lateral distance; S.Ed: standard error difference; CD: critical difference. *significance level at 0.05%; *significance at 0.01%; NS: not significant.
The various genotypes of rice had varying RLD values: 1.513cm/cm3 (JKRH 3333); 1.267 cm/cm3 (TNRH 180); and 1.077cm/cm3 [ADT(R)45]. Conventional aerobic rice observed a decreased RMD value of 1.133cm/cm3. Among the genotypes, JKRH 3333 had a higher RMD value (1.214mg/cm3) with statistical significance over TNRH 180 (1.109 mg/cm3) and ADT(R)45 (0.996 mg/cm3). The root density changed in the drip system, which was higher for the JKRH 3333 genotype than the other genotypes (Figure 1). The root dry weight (2.56 g/hill) and specific root length (0.160) was found higher in JKRH 3333 over the rest (Figure 2). Comparing the drip treatments, increased root dry weight observed in 0.8 m LD (2.5g/hill) laid out at 5–10 cm SDI over conventional rice (1.9g/hill).
The root ATPase activity of JKRH 3333 (33.1µg Pi/g/h) showed was more statistically significant supremacy than TNRH 180 (29.5µg Pi/g/h) and ADT(R)45 (23.8µg Pi/g/h). Among the drip irrigation treatments, increased root activity was obvious in 0.8m LD in SDI laid at the soil depth of 5–10cm treatment (31.2µg Pi/g/h) and lesser activity was evident in conventional aerobic rice (26.4µg Pi/g/h).
Genotypic variation of rice showed an increased filled grain percentage in JKRH 3333 (88.2%) followed by TNRH 180 (85.0%) and ADT(R) 45 (76.4%). The SDI system at 0.8m lateral distance was found to be higher (85.4%). Higher HI values were observed in JKRH 3333 (39.2%) followed by TNRH 180 (38.5%) and ADT(R) 45 (37.8%). The solar operated drip irrigation treatments were significantly superior with an elite value of 39.3% compared with the well-operated drip irrigation system (37.7%).
JKRH 3333 genotype was statistically superior among all the genotypes in grain yield. The grain yield was observed to be significantly higher in the solar operated drip irrigation treatment (4817kg/ha) compared with well-operated drip irrigation (4313kg/ha) (Table 2). Among the performance of genotypes under drip irrigated aerobic rice, JKRH 3333 was statistical superior in mean grain yield (4831kg/ha) followed by TNRH 180 (4639kg/ha) and ADT(R)45 (4224kg/ha).
LD: lateral distance; S.Ed: standard error difference; CD: critical difference. **significance level at 0.05%; *significance at 0.01%; NS: not significant.
Grain yield (kg/ha) | Source of drip irrigation | Varieties | Drip treatments | |||||||
---|---|---|---|---|---|---|---|---|---|---|
Solar | Well | TNRH 180 | JK RH 3333 | ADT (R) 45 | 0.8m LD | 1.2m LD | 5–10 cm | 15–20 cm | Conventional | |
4817 | 4313 | 4639 | 4831 | 4224 | 5121 | 4874 | 4302 | 4106 | 4421 |
Grain yield | Source of drip irrigation | Varieties | Drip treatments | |||
---|---|---|---|---|---|---|
S.Ed. | CD <0.05 | S.Ed. | CD <0.05 | S.Ed. | CD <0.05 | |
8.54 | 36.75** | 20.83 | 47.90** | 35.34 | 71.03** |
Roots are the main component in the absorption of water and minerals, which are essential in plant physiological processes. Fageria (2007) observed that root length followed a significant quadratic response with the advancement of plant age from 19 to 120 days after sowing, and shows a linear increase in root length during flowering.
Favored root length under SDI at 5–10cm treatment is due to deep rooting of rice to combat water limited conditions. Genotypic variation in TNRH 180 revealed deep rooting to reduce the limited water application effect. The increased root growth and development of the root system help the rice to explore the wider area of soil and the deeper soil layers for water and nutrients. These results were corroborated with Henry et al. (2011) in rice under drought.
The genotype JKRH 3333 registered an increased root length and specific root length of 34.9% and 3.9% over conventional aerobic rice (Figure 2). Specific root length was an indicator for environmental changes. The genetic potential of this rice genotype for maintenance of increased root length favors lateral root branching (Figure 3). This effect was in accordance with Kato & Okami (2011) in rice.
Root volume of plants covers huge soil volumes and water uptake from the soil in water-limited conditions (Kanbar, 2004). Altered root volumes were observed in the present study under SDI with a 5–10cm drip laid out at 0.8m LD, due to greater assimilation allocation in rice roots by drip irrigation. Similar results were observed by Parthasarathi et al. (2012) under drip irrigation.
The root length density (RLD) is the length of roots per unit volume of soil, is an important parameter required to understand plant performance. In the present experiment, the SDI at 5–10cm depth using JKRH 3333 increased the RLD and RMD, due to the root zone of rice exposed to frequent wetting and slight drying and nutrient accessibility. The dry weight of roots was 36.8% superior in JKRH 3333 hybrid under drip irrigation. Similar variation obtained in rice was observed by Vanitha (2011) and could support the present results. This unique response of root length and mass density under drip irrigation to improve nutrient and water accessibility was due to more root proliferation at topsoil. Comparing the root images of genotypes (Figure 3) revealed that, even though the appearance of white roots was common, an increase in root numbers and density was higher in drip irrigation.
Light energy absorbed by chlorophyll is converted into stable chemical energy and drives ATP formation via ATPase in the plastids of roots. ATPase is widely present in plant tissues and involved in the active transport of ions across membranes of the cell (Martínez-Ballesta et al., 2003). In the present study, higher levels of ATPases were observed in SDI + 0.8m LD at 5–10cm lateral depth with the JKRH 3333 hybrid.
The grain filling percentage is an important contributory factor to grain yield. The SDI laid out at 5–10cm depth with 0.8m LD treatment registered more grain production and filling percentage. Among the genotypes, the hybrid (JKRH 3333) excelled the variety in filled grain percentage by 15.4% (Figure 4). The increase in the water supply to the spikelets might reduce the floret abortion during flowering, and may be the reason behind higher filled grains in SDI. These results are indirectly supported by Kato et al. (2008) in aerobic rice.
Harvest index (HI) reflects the proportion of assimilate distribution between economic and total biomass (Donald & Hamblin, 1976). Among the genotypes, a higher in HI was recorded in JKRH 3333 with a 1.6 and 4.5% increment over the TNRH 180 and ADT(R)45 genotypes, respectively (Figure 3). This might be attributed to the fact that producing a larger sink size and efficient transport of assimilates from leaves and stems (‘source’) into developing spikelets (‘sinks’), thus resulting in the increased grain yield (Guan et al., 2010).
The higher grain yield of JKRH 3333 recorded a 21.4% increase in drip over conventional aerobic rice cultivation. Comparing the depth of SDI treatment at a 5–10cm soil depth achieved a 18.9 and 13.0% increased yield over 15–20cm soil depth and conventional aerobic irrigation method, respectively. The SDI system maintained equal soil wetting, reduced the evaporation with direct point application of water in root, which improves the grain yield of rice. A previous study supports this argument (Douh et al., 2013).
This drip-irrigated aerobic rice study concluded that there is an increase in grain yield along with increased root parameters. Based on the data of lateral spacing, discharge variations and the root characters of rice under drip significantly showed that there was characteristic flexibility in the roots of the rice plant. The root length, root density, root hairs and root ATPase activity exhibited a significant association with filled grain percentage and grain yield. The genotype JKRH 3333 showed 14.3% increased grain yield with favorable root density and root dry weight over ADT(R)45. It could be recommended that 0.8m lateral distance laid out at 5–10cm depth SDI may proliferated more roots at subsurface soil layer with a yield increment in rice.
Dataset 1: Response of root traits phenotyped under different drip irrigation treatments. doi, 10.5256/f1000research.9938.d151043 (Parthasarathi et al., 2017).
Source of irrigation: S1, solar powered; S2, well operated.
Drip treatments: T1, 0.8m LD; T2, 1.2m LD; T3, 5–10cm; T4, 15–20cm; T5, conventional aerobic rice.
Genotypes: V1, TNRH 180; V2, JKRH 3333; V3, ADT(R)45.
TP, SM, EV, and KV designed the experiments. TP performed the experiments in the field. TP and VM analyzed the data using statistics. TP, VM, and KV contributed reagents/materials/analysis tools. TP wrote the manuscript. KV and VM corrected the manuscript.
The project was funded by Netafim Irrigation Ltd., Israel.
The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Supplementary File 1: Detailed information on experimental set up (Table S1), genotypes (Table S2) and fertigation schedule (Table S3) of drip irrigated rice study.
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Competing Interests: No competing interests were disclosed.
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