Shoot Traits Research Articles

The use of seedling root and shoot traits to predict the saleable yield of elite red papaya genotypes

ABSTRACT Determining the saleable fruit yield of red hermaphrodite papaya (Carica papaya L.) relies on waiting until tree maturity. Recombinant inbred lines (RIL) seedlings (F6-F7 RIL) were assessed for fresh root length, fresh shoot weight, fresh shoot width, dry root weight and dry shoot weight during three growth stages after sowing (0-30 days after sowing; DAS, 31–60 DAS and 61–90 DAS). During the 31–60 DAS period, significant intra-genotypic variation was observed for each trait across the RIL assessed. A strong genotypic correlation at growth stage II was predicted among the traits (P < 0.05; rg > 0.60), with a moderate to high broad-sense heritability (H2 b.s; > 0.30) and with genetic advances of up to 10% in the RIL compared to the parental genotypes. Growth stage II was most reliable for predicting saleable fruit yield (R 2 = 0.73) with levels of influence of traits measured across the RIL in the following order: fresh root length (40% to 70%) > dry root weight (20% to 50%) > dry shoot weight (10% to 48%) > fresh shoot width (13% to 38%) > fresh shoot weight (1% to 42%). Hence, these F7 genotypes will be used to develop new F1 hybrids with significant increases in saleable fruit yield.

Screening and Identification of Aluminium Toxicity Tolerant Lentil (Lens culinaris Medik.) RILs based on Root Growth Studies and Organic Acid Exudation under Hydroponics

Background: Aluminium (Al) stress is among the prime limitations of crop production including Lentil, in acidic soils as Al solubilizes into phytotoxic forms at low pH causing root growth inhibition, minimizing plant-vigor and yield. The current study was performed to identify the Al tolerant RILs based on root-regrowth studies, short term growth culture and organic acid exudation in reaction to Al toxicity under hydroponics. Methods: The study involved screening for Al toxicity tolerance in recombinant inbred lines (RILs) population of lentil through root re-growth studies, short term growth response method under hydroponics treated with toxic concentration of Al (148 µM), organic acid exudation using HPLC and correlation analysis among the traits. Result: ANOVA for root and shoot traits revealed presence of highly significant genotypic differences for all the traits. High GCV along with high H2bs and GA% were observed for the traits revealing the influence of additive genes and suggested reliable selection of these traits for Al toxicity tolerance. Citric acid was exudated in highest amount in all the genotypes and was positively and significantly correlated with RRG. Based on the hydroponics study and organic acid exudation, RILs identified as tolerant were LRIL-10, LRIL-37, LRIL-68, LRIL-96, LRIL-97, LRIL-113, LRIL-125, LRIL-133, LRIL-143, LRIL-144 and LRIL-148. With further evaluation, these RILs may serve as important Al toxicity tolerant varieties suitable for acidic soil conditions. Also, these lines may serve as donors of Al toxicity loci and mapping of Al tolerance genes.

Genetic diversity, relationships among traits and selection of tropical maize inbred lines for low-P tolerance based on root and shoot traits at seedling stage.

The tropical maize breeding for low-P tolerance and good performance under low-P stress environments can be achieved through selection based on root morphology traits at seedling stage. Here, we assessed the genotypic variation and genetic diversity of a panel of 151 tropical maize inbred lines for root and shoot seedling traits, investigated the relationship among traits and selected a set of promising inbred lines for low-P tolerance and performance. We evaluated the inbred lines at seedling stage in a greenhouse experiment under two conditions: applied P (AP) and non-applied P (NAP). A mixed model approach was used to estimate variance components and predict the genotypic values of each inbred line. The genetic diversity among inbred lines based on root and shoot traits was assessed, and correlations were estimated between tested traits under AP and NAP. Our panel of inbred lines showed huge genetic variability for all traits and presented large genetic diversity under both P conditions. Variance components due to the inbred line × P condition interaction were also highly significant (P < 0.01) for all traits. Root dry weight (RDW) was positively associated with stalk dimeter (SD), shoot dry weight (SDW) and root length, volume, and area under both P conditions. Also, the SD and SDW were associated with most root traits under AP. Based on low-P tolerance and performance indices, we selected a set of top 20 inbred lines to be used in our maize breeding program. We therefore concluded that there is a significant genetic diversity in the tropical maize inbred lines which have the genetic potential to be use in association mapping studies and also to develop improved low-P tolerant and P-efficient hybrids and maize breeding populations for low-P stress environments.

Salt tolerance in wheat is associated with the maintenance of shoot biomass, stomatal conductance, and sucrose in the phloem.

Wheat (Triticum aestivum L.) is a mega-staple for millions of the world's populations and its yield potential is impacted by soil salinization. This study investigated genotypic variation in salt tolerance among six wheat genotypes, Gladius, Drysdale, GD0014, GD0120, GD0180, and GD0185. The study also characterized shoot traits, photosynthetic traits, leaf Na and K concentrations, and phloem sucrose. The plants were grown under controlled growth room conditions at 0 mM NaCl (Control) and 100 mM NaCl. The results showed that the salt tolerance index (STISFW, SFW: shoot fresh weight) varied from 0.52 for GD0120 to 0.69 for GD0180. Based on the STISFW, salt tolerance for the wheat genotypes was in the order, GD0180 > Gladius > GD0185 > Drysdale > GD0014 > GD0120. Projected shoot area (PSA) at all growth stages, 14, 20, 27, 34, and 40 DAS were strongly correlated with SFW at 45 DAS. Salt treatment significantly increased phloem sucrose level in the salt intolerant, Drysdale, while having no effect on this parameter in Gladius. Gladius showed greater maintenance of stomatal conductance than Drysdale. The relative ratio of K/Na between treatment and control was strongly correlated with the relative ratio of SFW (r = .85). The correlation between PSA at 14 DAS and SFW at 45 DAS and the correlation between the relative ratio of K/Na between treatment and control with STISFW identify these parameters to be potential traits for screening salt tolerance in wheat. Higher salt tolerance in Gladius would be associated with higher maintenance of stomatal conductance and enhanced phloem sucrose transport.

Comprehensive phenotyping of SKUAST-K released rice varieties reveals significant role of root traits in drought resilience

The North Western Himalayan region, particularly the Kashmir valley, harbors a rich diversity of rice landraces, yet systematic studies on climate resilience are lacking. The present study was aimed at investigating the root traits and their interplay with shoot morphological and physiological traits under drought stress using seven indica and three japonica rice varieties released by SKUAST-Kashmir and their possible role in adaptive capacity to different environments. Significant variability was observed in root and shoot traits across the varieties. Root traits exhibited a wide phenotypic range, including rooting depth, fresh and dry weight, diameter, length, volume, surface area, and length density. Associations between root and shoot traits were identified, with notable correlations such as shoot weight with root shoot ratio, shoot length with root dry weight, diameter, and surface area, and tiller number with various root traits. Additionally, physiological traits like canopy temperature depression showed significant correlations with root depth and surface area. Principal component analysis (PCA) analysis grouped japonica type varieties together with a notable outlier, Chenab, due to its superior root traits among indica varieties. These findings emphasize the substantial contribution of root traits to productivity and advocate for their integration into varietal development processes. In view of the increasing evidences in crop plants about their role of root traits in defining plant productivity and reproductive fitness, study of root traits could provide valuable insights into the patterns of crop adaptation to diverse areas and growing environments.

Combination of Polymer-Coated Urea and Rapid-Release Urea Increases Grain Yield and Nitrogen Use Efficiency of Rice by Improving Root and Shoot Activities

The use of polymer-coated urea (PCU) can improve nitrogen use efficiency (NUE), compared to the application of rapid-release urea (RU). However, the effect of PCU-based nitrogen management on grain yield and the NUE of rice and its underlying mechanism remain unclear. A japonica rice cultivar Jinxiangyu 1 was grown in the field with four treatments including N omission (0N), split application of RU (Control), one-time application of 100% PCU (T1), and one-time application of 70% PCU + 30% RU (T2). Results showed that, compared to the control, the grain yield was significantly increased in the T2 treatment, while it was comparable in the T1 treatment. This was mainly due to increased total spikelets in the T2 treatment. Root oxidation activity (ROA) and root zeatin (Z) + zeatin riboside (ZR) content during booting were the distinct advantages of the T2 treatment, compared to either the control or T1 treatment, exhibiting significant or highly significant correlations with leaf photosynthesis. This process contributed significantly to total spikelets and total N uptake. Additionally, the T2 treatment absorbed more N than the control without reducing the internal N use efficiency (IEN), primarily due to its unchanged harvest index (HI) driven by comparable non-structural carbohydrate remobilization. In conclusion, combining PCU with RU can enhance the coordination of root and shoot traits during booting while maintaining a competitive HI at maturity, thereby significantly improving grain yield and achieving a balance in N uptake and utilization.

Multigenerational invasive plant competition causes greater root than shoot trait shifts in a perennial grass

Multigenerational invasive plant competition causes greater root than shoot trait shifts in a perennial grass

Differential spatial plasticity response in common bean (Phaseolus vulgaris L.) root architecture under water stress is driven by increased root diameter, surface area and volume at deeper layers

Root plasticity enables plants to adapt to spatial and temporal changes in soil resources. In this study, 40 common bean genotypes evaluated for two root and shoot traits under irrigated and water stress. Three genotypes WB-216, WB-N-2, and WB-966 with contrasting plasticity responses were used for in-depth study. Highest positive plasticity for most root traits was found in case of WB-N2 and WB-216, whereas, WB-966 had negative plasticity for all the traits recorded. In terms of spatial plasticity for root traits in three root length sections, WB-216 was positively plastic for root diameter with progressive decrease from top to bottom sections. WB-N2 had positive plasticity values for root diameter, root surface area and root volume. WB-966 had negative plasticity for all the traits. For WB-216, the root diameter increased under drought in S1 but was almost same in bottom sections. In case of WB-N2, there was increase in root diameter in S2 and S3, but for WB-966, root diameter decreased in all sections. Similar trend was observed in all three genotypes for root surface area and volume. We report that major drivers of spatial plasticity of root architectural traits are increased root diameter, surface area and volume at deeper layers.

Multi-locus genome-wide association analysis for root and shoot traits at seedling stage in Ethiopian sorghum (Sorghum bicolor (L.) Moench) accessions

Multi-locus genome-wide association analysis for root and shoot traits at seedling stage in Ethiopian sorghum (Sorghum bicolor (L.) Moench) accessions

Common‐garden study of introgression at loci associated with traits adaptive to coastal environment from Quercus dentata into Quercus mongolica var. crispula

AbstractAdaptive introgression, which can provide genetic diversity resulting in the expansion to novel habitats and the creation of different ecotypes, has been proposed in oaks (genus Quercus). In northern Japan, Quercus mongolica var. crispula (Qc) is common in inland habitats, and Quercus dentata (Qd) occurs in coastal habitats. At the northern distributional limit of Qd, Quercus × angustilepidota (Qa), a hybrid taxon between Qc and Qd, occurs in coastal habitats. These taxa, Qc, Qa, and Qd, were transplanted to inland and coastal common gardens (sites). Genotypes at 27 495 loci, phenotypes of eight leaf and shoot traits, and the size of 30‐year‐old trees were measured for 224 individuals in both sites. Genotypic variation revealed a hybrid zone between Qc and Qd, including both northern‐edge Qd admixed with Qc and coastal Qa backcrossed to Qc. Phenotypes of Qa trees were intermediate between those of Qc and Qd trees. The size of Qa and Qd trees was smaller than that of Qc trees in the inland site but was larger in the coastal site, suggesting adaptation of Qa and Qd to coastal environment. Local ancestry was estimated from phased genotypes of admixed trees using reference genotypes of 47 Qc and 25 Qd trees, indicating heterogeneous ancestry along chromosomes. Admixture mapping of local ancestry detected loci significantly associated with a leaf trait, which were located in an introgressed genomic region with genes related to stress response. Further studies are necessary to elucidate the genetic basis of adaptive introgression resulting in Qd‐like phenotypes of Qa.

Variability of Root and Shoot Traits under PEG-Induced Drought Stress at an Early Vegetative Growth Stage of Soybean

Variability of Root and Shoot Traits under PEG-Induced Drought Stress at an Early Vegetative Growth Stage of Soybean

Nutritional Variation on Sequentially Harvested Shoots and Fruits of Lagenaria siceraria Landraces.

Lagenaria siceraria (Molina) Standley, a member of the Cucurbitaceae family, is valued for its medicinal and nutritive properties. The nutrient status of sequentially harvested shoots and fruits at various growth stages in L. siceraria has not been documented to date. This study aimed to compare the nutritional status of L. siceraria landrace shoots and fruits harvested at different maturity stages. Micronutrients, macronutrients, and proximate composition of shoots and fruits were determined using inductively coupled plasma-optical emission spectrometry (ICP-OES). Data were subjected to multivariate statistical analysis. The nutrient attributes differed significantly (p < 0.05) within and among landraces at different growth stages. Correlation of nutritional traits was primarily based on shared absorption sites and comparable chemical composition. The first five principal components in shoots and fruits had 90.218 and 89.918% total variability, respectively. The micronutrients Ca, Mg, K, P, and N in shoots and the macronutrients Fe, Zn, Cu, and Al in fruits were the main contributors to variability. The biplot and dendrogram clustered landraces with comparable nutrient values. Shoot traits classified landraces into three major clusters, where clusters I and II grouped landraces with superior and inferior Mg, P, K, N, Fe, K/Ca+Mg, ADL, and protein levels at 42-63 DAS. Cluster III consisted of landraces with more Ca, Na, Mn, Zn, and Cu. However, the fruit nutrient status classified landraces into two major clusters. Cluster I comprised landrace KSC (as a singlet) with the highest Ca, P, N, Mn, Fe, Zn, and Cu contents at various stages of growth (7-21 DAA). Cluster II had landraces with higher K, C/N ratio, Na, moisture content, ash, protein, and fat. The nutritional status of shoots and fruits determined at different stages of growth is essential for selecting the best harvest time and landrace(s) for required daily nutrient intake.

Variations in water economy traits in two Sphagnum species across their distribution boundaries.

We assessed changes in traits associated with water economy across climatic gradients in the ecologically similar peat mosses Sphagnum cuspidatum and Sphagnum lindbergii. These species have parapatric distributions in Europe and have similar niches in bogs. Sphagnum species of bogs are closely related, with a large degree of microhabitat niche overlap between many species that can be functionally very similar. Despite this, ecologically similar species do have different distributional ranges along climatic gradients that partly overlap. These gradients may favor particular Sphagnum traits, especially in relation to water economy, which can be hypothesized to drive species divergence by character displacement. We investigated traits relevant for water economy of two parapatric bryophytes (Sphagnum cuspidatum and S. lindbergii) across the border of their distributional limits. We included both shoot traits and canopy traits, i.e., collective traits of the moss surface, quantified by photogrammetry. The two species are ecologically similar and occur at similar positions along the hydrological gradient in bogs. The biggest differences between the species were expressed in the variations of their canopy surfaces, particularly surface roughness and in the responses of important traits such as capitulum mass to climate. We did not find support for character displacement, because traits were not more dissimilar in sympatric than in allopatric populations. Our results suggest that parapatry within Sphagnum can be understood from just a few climatic variables and that climatic factors are stronger drivers than competition behind trait variation within these species of Sphagnum.

From buds to shoots: insights into grapevine development from the Witch's Broom bud sport.

Bud sports occur spontaneously in plants when new growth exhibits a distinct phenotype from the rest of the parent plant. The Witch's Broom bud sport occurs occasionally in various grapevine (Vitis vinifera) varieties and displays a suite of developmental defects, including dwarf features and reduced fertility. While it is highly detrimental for grapevine growers, it also serves as a useful tool for studying grapevine development. We used the Witch's Broom bud sport in grapevine to understand the developmental trajectories of the bud sports, as well as the potential genetic basis. We analyzed the phenotypes of two independent cases of the Witch's Broom bud sport, in the Dakapo and Merlot varieties of grapevine, alongside wild type counterparts. To do so, we quantified various shoot traits, performed 3D X-ray Computed Tomography on dormant buds, and landmarked leaves from the samples. We also performed Illumina and Oxford Nanopore sequencing on the samples and called genetic variants using these sequencing datasets. The Dakapo and Merlot cases of Witch's Broom displayed severe developmental defects, with no fruit/clusters formed and dwarf vegetative features. However, the Dakapo and Merlot cases of Witch's Broom studied were also phenotypically different from one another, with distinct differences in bud and leaf development. We identified 968-974 unique genetic mutations in our two Witch's Broom cases that are potential causal variants of the bud sports. Examining gene function and validating these genetic candidates through PCR and Sanger-sequencing revealed one strong candidate mutation in Merlot Witch's Broom impacting the gene GSVIVG01008260001. The Witch's Broom bud sports in both varieties studied had dwarf phenotypes, but the two instances studied were also vastly different from one another and likely have distinct genetic bases. Future work on Witch's Broom bud sports in grapevine could provide more insight into development and the genetic pathways involved in grapevine.

Water Use Strategies and Shoot and Root Traits of High-Yielding Winter Wheat Cultivars under Different Water Supply Conditions

Drought is the most important factor limiting winter wheat yield in the North China Plain (NCP). Choosing high-yielding cultivars is an important measure to minimize the negative effects of drought stress. Field studies were conducted with 10 cultivars in the 2020–2022 seasons under three irrigation treatments (I0, without irrigation; I1, irrigated at jointing stage; I2, irrigated at jointing and anthesis stages) in the NCP to examine the water use strategies and root and shoot traits of high-yielding cultivars under different water supply conditions. The results showed that yield variation among cultivars was 21.2–24.6%, 23.7–25.9% and 11.6–15.3% for the I0, I1 and I2 treatments, respectively. Under water deficit conditions (I0 and I1), high-yielding cultivars reduced water use during vegetative stages and increased soil water use during reproductive stages, especially water use from deeper soil layers. Those cultivars with higher root length density (RLD) in deep soil layers exhibited higher water uptake. Each additional millimeter of water used after anthesis from the 100–200 cm soil layers increased grain yield by 23.6–29.6 kg/ha and 16.4–28.5 kg/ha under I0 and I1, respectively. This water use strategy enhanced dry matter accumulation after anthesis, decreased canopy temperature (CT) and increased relative leaf water contents (RLWC), which ultimately improved grain yield. For winter wheat grown under I2, cultivars that decreased water use after anthesis had higher water productivity (WP). Root length (RL), root weight (RW) and root:shoot ratio were each negatively correlated with grain yield, while above-ground biomass was positively correlated with grain yield. Therefore, higher dry matter accumulation and smaller root systems are two important traits of high-yielding cultivars under sufficient water supply conditions (I2) in the NCP.

Combined Vinasse and Mineral NPK Fertilizer Affect Physio-Biochemical, Root, and Yield Characters of Faba Bean (vicia faba l.) Genotypes Grown on Saline Soil

The objective of this study is to assess the effects of combined applications of recommended traditional chemical fertilization NPK 100% without Vinasse and with 100, 75, 50 NPK + Vinasse on Physio-Biochemical, root, shoot traits, yield, and its attributes for four faba bean Giza-429, Giza-843, Misr-2 and Sakha-3 genotypes which are cultivated in the soil is salt affected of eastern Egypt. The experimental design was conducted using a strip-plot design with three replicates in 2020 and 2021. Treatments of different doses of chemical fertilization were 100% NPK without Vinasse and 100%, 75%, and 50% of NPK with Vinasse as supplementary organic fertilizer were arranged in vertical strips, while horizontal strips were devoted to faba bean genotypes (Vicia faba L.) i.e. Giza-429, Giza-843, Misr-2, and Sakha-3. Relative to the other fertilizer combinations, the 75% NPK + Vinasse did not contribute more than 5% more yield to Sakha 3 than it did with the 100% NPK + Vinasse. Both the fertilizer and cultivar main effects had the same positive linear trend, with relatively higher total seed yield in the case of the 75% NPK + Vinasse and of cv. Sakha 3. The harvest index (HI) was different only for fertilizer levels.The 75% NPK + Vinasse differed by a very negligible margin from the 100% NPK + Vinasse, but both differed from the other two levels. Along with seed yield, both straw and biological yields were significant for the interaction effects, as well as for the main effects. Within all fertilizer combinations, cultivars Giza 843 and Saka 3 consistently, as a subgroup, outyielded the other two cultivars, not only for total seed yield but also for all other yield component characters.

Drought effects on shoot traits and introduction of new indices in chickpea to identify drought tolerant and susceptible genotypes

Drought effects on shoot traits and introduction of new indices in chickpea to identify drought tolerant and susceptible genotypes

Impact of ultraviolet-B radiation on early-season morpho-physiological traits of indica and japonica rice genotypes.

Ultraviolet (UV)-B radiation is considered one of the major detrimental rays coming from the Sun. UV-B radiation has a harmful impact on plant growth and development. The effect of UV-B radiation was studied on 64 rice (Oryza sativa L.) genotypes during the vegetative season. An equal number of genotypes from the japonica (50%) and indica (50%) subspecies were phenotyped using the Soil-Plant-Atmosphere-Research (SPAR) units. The 10 kJ UV-B was imposed 12 days after planting (DAP) and continued for three weeks (21 d). Based on the combined ultraviolet-B radiation response index (CUVBRI) for each genotype, the 64 rice genotypes were classified into sensitive, moderately sensitive, moderately tolerant, and tolerant. Various shoot traits, such as plant height, tiller, and leaf numbers, were measured. We also studied critical root phenological traits like root volume, diameter, tips, and forks. Out of all the studied shoot traits, leaf area showed maximum reduction for both indica (54%) and japonica (48%). Among the root traits, root length decreased by negligible (1%) for indica as compared to japonica (5%), while root crossing and forks showed a maximum decline for japonica (37 and 42%), respectively. This study is timely, meaningful, and required because it will help breeders select a tolerant or sensitive rice line for better yield and production under abiotic stresses.

Temporal phenotypic variation of spinach root traits and its relation to shoot performance

The root system is important for the growth and development of spinach. To reveal the temporal variability of the spinach root system, root traits of 40 spinach accessions were measured at three imaging times (20, 30, and 43 days after transplanting) in this study using a non-destructive and non-invasive root analysis system. Results showed that five root traits were reliably measured by this system (RootViz FS), and two of which were highly correlated with manually measured traits. Root traits had higher variations than shoot traits among spinach accessions, and the trait of mean growth rate of total root length had the largest coefficients of variation across the three imaging times. During the early stage, only tap root length was weakly correlated with shoot traits (plant height, leaf width, and object area (equivalent to plant surface area)), whereas in the third imaging, root fresh weight, total root length, and root area were strongly correlated with shoot biomass-related traits. Five root traits (total root length, tap root length, total root area, root tissue density, and maximal root width) showed high variations with coefficients of variation values (CV ≥ 0.3, except maximal root width) and high heritability (H2 > 0.6) among the three stages. The 40 spinach accessions were classified into five subgroups with different growth dynamics of the primary and lateral roots by cluster analysis. Our results demonstrated the potential of in-situ phenotyping to assess dynamic root growth in spinach and provide new perspectives for biomass breeding based on root system ideotypes.

Agronomic and Physiological Performance of the Indica Rice Varieties Differing in Tolerance to Low Phosphorus

Phosphorus (P) deficiency and low P use efficiency (PUE) are limiting factors in rice (Oryza sativa L.) production. Understanding the agronomic and physiological traits of P-tolerant rice varieties is crucial for improving PUE. However, the agronomic and physiological traits of rice varieties differing in tolerance to low P have not been fully studied or comprehensively explored. Two varieties with strong tolerance to low P (STVs, low P tolerance index &gt; 0.9) and two with weak tolerance to Low P (WTVs, low P tolerance index &lt; 0.5) were grown hydroponically with normal P level (NP, 8.02 mg L−1) and low P level (LP, 0.401 mg L−1) in year 2020 and 2021. Results showed that, compared with NP, the LP significantly decreased grain yield, but enhanced P translocation efficiency (PTE), internal P use efficiency (IPE), and P harvest index (PHI) in all the varieties. The STVs showed better performance than the WTVs. Specifically, the STVs exhibited a 131.33% higher grain yield, 15.95% higher PTE, 41.6% higher IPE, and 8.84% higher PHI compared to the WTVs. The STVs also exhibited superior shoot traits, including increased productive tillers, leaf area index (LAI), leaf photosynthetic rate, shoot biomass, contents of indole-3-acetic acid (IAA) and zeatin (Z) and zeatin riboside (ZR) in leaves, non-structural carbohydrates (NSC) remobilization during grain filling, and content of NSC per spikelet, when compared to the WTVs under the LP treatment. Additionally, the STVs demonstrated better root traits, such as higher root biomass, root oxidative activity (ROA), root acid phosphatase (RAP) activity, and greater root IAA and Z + ZR contents. These shoot and root traits exhibited highly positive correlations with grain yield, PTE, and IPE. In conclusion, the STVs maintain higher grain yield and PUE under the LP treatment, due mainly to their improved root and shoot agronomic and physiological traits, which provide valuable references for selecting for P-efficient rice varieties.