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Climatic Conditions and Production of Soybean in Northeastern Brazil
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This genotype was also ranked as one among the top three for total root volume, fine root length, and fine root surface area. Similarly, genotype NMS exotic pedigree was ranked as one among the top three for total root length, total root surface area, total root volume, fine root length, and fine root surface area, and as one among the top five for average root diameter. Another genotype with exotic pedigree, N, was ranked as one among the top 10 for total root length, total root surface area, total root volume, fine root length, and fine root surface area.
Genotype PI G. This genotype was also ranked as one among the lowest 10 for fine root length and fine root surface area. Genotype RF sustained nitrogen fixation under drought conditions was ranked as one among the lowest 10 for total root length, total root surface area, total root volume, average root diameter, fine root length, and fine root surface area. We conducted a principal component analysis PCA based on all phenotypic data and generated a biplot to investigate the possibility of clustering of genotypes Fig 3.
The biplot separated the genotypes in to seven clusters.
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Genotype PI , which had the lowest total root length, total root surface area, total root volume, and average root diameter, was clearly separated from all other genotypes Cluster 7. All genotypes that were ranked among the top 10 for at least three root traits Clusters 1, 2, and 3 were included in the quadrants 1 and 4, whereas, all genotypes that were ranked among the bottom 10 for at least three root traits Clusters 4, 5, 6, and 7 were included in the quadrants 2 and 3.
The most important root traits contributing to the clustering pattern were total root surface area, total root length, total root volume, fine root length, and fine root surface area. Genotypes 1—49 are marked on the biplot; please see Table 1 for the genotype names corresponding to the numbers. Plant height was not related with total root length, fine root length, and fine root surface area, and was negatively correlated with total root surface area r, Seed size did not have any significant relation with total root length, total root surface area, total root volume, average root diameter, fine root length, and fine root surface area Table 5.
Fine root traits were positively correlated with whole root system traits Table 5. Considerable variability was detected for root traits in the soybean germplasm collection of 49 genotypes evaluated in this study. These genotypes were selected based on a variety of traits that are important for soybean improvement e.
The variability of root traits we identified among the 49 genotypes is promising and warrants additional research to further explore the genetic diversity in wild and domesticated soybean. The extent of variability for root traits among the soybean genotypes is demonstrated by the wide range observed for these traits Table 2. However, maturity groups did not influence any root traits [P-values for the effect of maturity groups on total root length, total root surface area, total root volume, average root diameter, fine root length, and fine root surface area were 0.
Similar observations were made by Turman et al. This study evaluated root penetration ability of soybean genotypes using wax-petroleum jelly discs, which simulate compacted soil layers or soil hardpans. Analysis of variance detected significant interaction between run and genotype for root penetration Table 2 , and we analyzed the penetration data separately for each run Table 3. Temperature influences the penetration resistance of the wax- petroleum jelly hardpans S1 Fig.
The differences in weather conditions during Run 1 and 2 might have influenced the greenhouse temperature slightly, which in turn influenced the penetration resistance of the hardpans. This might be the reason for differences in root penetration of genotypes between runs. To the best of our knowledge, this study is the first one evaluating a diverse soybean germplasm collection for root penetration. Soil compaction occurs in nearly every farm in the United States, limiting root penetration and crop yields. In the southeastern United States, most soils have an inherent compacted layer of subsoil hardpan , which often necessitates expensive and non-sustainable tillage operations to increase the rooting zone.
Our study has identified soybean genotypes that penetrated the synthetic hardpans Table 3. We found that eighteen genotypes penetrated the hardpan fully or partially in at least one run, and the behavior was consistent in both runs for six of them NC-Raleigh, N, N, LG, Benning, and Crockett. These genotypes offer useful genetic material for breeders to develop high yielding soybean varieties for hardpan forming soils. We have presented 10 genotypes that were ranked high and 10 genotypes that were ranked low for total root length, surface area, and volume, average root diameter, and fine root length and surface area in Table 4.
Although the physiological basis for slow wilting is not yet determined, it likely involves root traits that improve water use efficiency or water conservation during soil drying [ 58 ]. Thus, it could be reasoned that the increased length, surface area, and volume of the whole root system and the fine roots contribute to the slow wilting ability of the genotype NTCPR The slow wilting nature of these genotypes combined with their ability to penetrate the hardpans makes them valuable genetic materials for breeding for drought tolerance in hardpan forming soils like that exists in the Southeastern United States.
In our study, we found that the fine root traits were related with the whole root system traits Table 5. Similar observations are reported by Prince et al. Fine roots increase root surface area per unit mass [ 60 ]. Since they are the most active part of the root system in extracting water and nutrients [ 61 , 62 , 63 ], the enhanced resource capture achieved through fine roots might have increased total root length, surface area, and volume as well. In the present research, shoot dry weight and chlorophyll index were positively correlated with total root length, total root surface area, total root volume, and fine root length Table 5 , S2 Fig.
Shoot dry weight and chlorophyll index are easily selectable traits, and are commonly utilized by soybean improvement programs to select desired genotypes. Since selecting genotypes based on root traits is highly challenging in a soybean breeding program, the positive correlations of shoot dry weight and chlorophyll index with root traits are advantageous as the genotypes selected based on these easily measurable shoot traits can have improved root systems as well.
Water and nutrient uptake from the soil is proportional to the contact area between root surface and soil [ 64 ]. This indicates that resource uptake increases with root surface area. Liang et al. Hudak and Patterson [ 65 ] found that a large root system, influenced by root length, surface area, and volume, enables the plant to exploit substantial soil volume, and is crucial for improving yield under drought conditions in soybean. In the present study, the increased resource capture achieved through larger root systems that were realized by increased root length, surface area, and volume might have contributed to increased dry matter addition, and thus, shoot dry weight.
Additionally, better nitrogen uptake achieved through larger root systems might have contributed to increased chlorophyll index. On the other hand, the increased amount of photoassimilates as a result of increased leaf greenness measured through chlorophyll index and shoot growth might have been utilized to increase root growth. Taken together, our results suggest that chlorophyll index and shoot weight have the potential to be indirect selection criteria for root traits that contribute to high yield potential.
The absence of correlation between plant height and total root length and the negative correlations of plant height with total root surface area and total root volume do not support the view that selecting for decreased plant height can result in a small root system. These results are supported by our own previous research along with that of others on multiple crops including chickpea [ 66 ], field pea Pisum sativum L. Total root length is determined by number and length of lateral roots [ 67 ], and is primarily controlled by different sets of genes, compared to plant height [ 68 ].
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The negative correlations of plant height with total root surface area and total root volume may be because assimilates that are not used to increase plant height might have diverted to root system to add more surface area, and thus, volume. Contrasting reports exist in terms of correlation of seed size with root traits [ 44 , 69 , 70 ].
Seed size was not correlated with any root traits evaluated in the present research Table 5. This shows that large seeds may not always produce long roots or large root systems. In the United States, soybean breeders have pursued the promising approach of introducing exotic germplasm to their breeding programs to increase genetic diversity. This approach has been found to be useful for improving yield and drought tolerance [ 57 , 58 , 71 ]. Twelve soybean lines with exotic pedigree, which were included in the South Carolina breeding program, were tested in the present study for root traits.
The soybean germplasm tested in this study included three G. Two of them PI and PI were ranked in the lowest 10 for most at least three root traits Table 4.
Our results are supported by previous reports that root and shoot growth of G. This variability should be considered when making interspecific hybridizations in breeding programs. Interestingly, genotype NMS, which was ranked as one among the top three for total root length, total root surface area, total root volume, fine root length, and fine root surface area, and as one among the top five for average root diameter, had G.
Significant genetic variability was observed for root traits in the soybean germplasm collection of 49 genotypes that was examined. Genotypes NTCPR slow wilting , NMS exotic pedigree , and N exotic pedigree were ranked high and genotypes PI wild and RF sustained nitrogen fixation under drought conditions were ranked low for most root traits. To our best knowledge, the present study is the first one evaluating a diverse soybean germplasm collection for root penetration. The genotypes that were able to penetrate the synthetic hardpan offer useful genetic material for breeding programs to improve yield in hardpan forming soils like that exists in the Southeastern United States.
We also examined whether root traits were related with plant height, shoot dry weight, chlorophyll index, and seed size, and found that only shoot dry weight and chlorophyll index were positively related with root traits, plant height was not correlated or had negative correlations with root traits, and seed size was not related with any root traits.
There were five replicated jars at each temperature. We thank Dr. Thomas E. Carter for consultation and for providing us with material that was used in this study, Dr. William Bridges and Dr. Pat Gerard for statistical consultation, and Dr. We also thank Mr. Ricardo St. Aime, Ms. Amanda Williams, Ms. Charlotte Snook, and Mr. Nicholas Accardo for their help in data collection.
This publication is Technical Contribution No.
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The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. National Center for Biotechnology Information , U. PLoS One.
Published online Jul Ricardo Aroca, Editor. Author information Article notes Copyright and License information Disclaimer. Competing Interests: The authors have declared that no competing interests exist. Received Feb 28; Accepted Jun This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. This article has been cited by other articles in PMC. Associated Data Supplementary Materials S1 Fig: Strength penetration resistance of wax-petroleum jelly mixture as a function of temperature.
S2 Fig: Relation of total root length, surface area, and volume with shoot dry weight, chlorophyll index, and plant height of soybean genotypes.
Environmental Stresses in Soybean Production : Mohammad Miransari :
S1 File: Excel file containing all data on root, shoot, and seed traits. Abstract Root systems that improve resource uptake and penetrate compacted soil hardpan are important for improving soybean Glycine max L. Introduction Soybean Glycine max L. Table 1 Soybean genotypes used in the study, their maturity group, and characteristics. Open in a separate window. Experimental details This research was conducted under controlled environmental conditions in a greenhouse at the Department of Plant and Environmental Sciences, Clemson University, Clemson, SC. Fig 1. The mesocosm used to grow soybean plants in the experiment.
Data collection Plant height and chlorophyll index were measured at the time of harvest. Statistical analyses The experimental design was a randomized complete block with four replications in both runs. Results Genetic variability of root traits Significant variability was observed for root traits among the soybean genotypes Table 2. Fig 2. The y-axis indicates the absolute number of genotypes in each root trait class. Table 2 Analysis of variance results on effects of run the study was conducted two times, which were designated as two runs , rep run , genotype, and run x genotype interaction and range for various root traits.
Table 3 Soybean root penetration of synthetic hardpans 2 cm thickness that simulate compacted soil layers. Fig 3. Principal component analysis biplot that separated the soybean genotypes in to clusters based on the root and shoot traits. Table 5 Correlations among various root and shoot traits of the 49 soybean genotypes.
Discussion Considerable variability was detected for root traits in the soybean germplasm collection of 49 genotypes evaluated in this study. Conclusions Significant genetic variability was observed for root traits in the soybean germplasm collection of 49 genotypes that was examined. Supporting information S1 Fig Strength penetration resistance of wax-petroleum jelly mixture as a function of temperature. TIF Click here for additional data file. S2 Fig Relation of total root length, surface area, and volume with shoot dry weight, chlorophyll index, and plant height of soybean genotypes.
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S1 File Excel file containing all data on root, shoot, and seed traits. XLSX Click here for additional data file. Acknowledgments We thank Dr. Data Availability All relevant data are within the paper and its Supporting Information files. References 1. SoyStats A reference guide to important soybean facts and figures. American Soybean Association. Soybean yield potential—a genetic and physiological perspective. Crop Sci. Soybeans: improvement, production, and uses agronomy monograph Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits.
J Exp Bot. Lynch J. Root architecture and plant productivity. Plant Physiol. Carter TE Jr. Breeding for drought tolerance in soybean: where do we stand? In: Pascale AJ, editor. World Soybean Conf. Poorter H, Nagel O. The role of biomass allocation in the growth response of plants to different levels of light, CO 2 , nutrients and water: a quantitative review. Aust J Plant Physiol. Fitter AH. Plant roots: the hidden half.
Genotypic variation in seedling root architectural traits and implications for drought adaptation in wheat Triticum aestivum L. Plant Soil. Improving drought-avoidance root traits in chickpea Cicer arietinum L. Plant Prod. Genetic variability for root morph-architecture traits and root growth dynamics as related to phosphorus efficiency in soybean.
Funct Plant Biol. QTL analysis of root traits as related to phosphorus efficiency in soybean. Ann Bot. Root growth of four common bean cultivars in relation to drought tolerance in environments with contrasting soil types. Exp Agric. Genetic variation in root traits and nutrient acquisition of lentil genotypes. J Plant Nutr. Mechanical impedance to root growth: a review of experimental techniques and root growth responses. Soil Sci. Effect of soil compaction on growth, yield and root rots of white beans in clay loam and sandy loam soil.
Soil Biol. Use of wax-petrolatum layers for screening rice root penetration. Effects of a compacted subsoil layer on root and shoot growth, water use and nutrient uptake of winter wheat. J Agric Sci.
Residual effects of tillage on Coastal Plain soil strength. Soil Science. Akpertey A. Genetic introgression from Glycine tomentella to soybean to increase seed yield. Germplasm Resources Information Network. Effect of flood stress on soybean seed germination in the field. Am J Plant Sci.
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