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The compositional nutrient diagnosis-CND method is a standard tool for evaluating plant nutritional status. Adjustments are crucial to elevate accuracy. The effectiveness of such methodological refinements should be rigorously assessed through accuracy tests that are benchmarked against the prescient diagnostic analysis-PDA methodology. The objective of this investigation was to refine the CND technique for a more precise evaluation of N, P, and B nutrient status in cotton. The study's database encompasses 144 data points pertaining to crop yield and foliar nutrient concentrations from cotton plantations in the Cerrado biome of Brazil. Subsequently, the CND norms were established through rigorous calibration. Three separate nutrient-dose trials, each featuring four levels of N, P and B, were carried out to assess plant true nutritional status. Adjustments were made to the nutrient responsiveness range-NRr (0.5 and 1.0), while yield response-YR were scrutinized at threshold levels (5% and 10%). The prerequisites for achieving high diagnostic accuracy were nutrient specific. For N, maximal accuracy was linked only to the YR parameter (YR = 10%). For P, the most precise outcomes were attained with a NRr = 0.5 and YI = 5%. For B, highest diagnostic accuracy when the NRr = 1.0 and YI = 10%. These insights highlight the need to fine-tune the CND method for reliable nutritional evaluations and cotton crop productivity optimization., (© 2024. The Author(s).)

Fertilization with nickel (Ni) can positively affect plant development due to the role of this micronutrient in nitrogen (N) metabolism, namely, through urease and NiFe-hydrogenase. Although the application of Ni is an emerging practice in modern agriculture, its effectiveness strongly depends on the chosen application method, making further research in this area essential. The individual and combined effects of different Ni application methods-seed treatment, leaf spraying and/or soil fertilization-were investigated in soybean plants under different edaphoclimatic conditions (field and greenhouse). Beneficial effects of the Soil, Soil + Leaf and Seed + Leaf treatments were observed, with gains of 7 to 20% in biological nitrogen fixation, 1.5-fold in ureides, 14% in shoot dry weight and yield increases of up to 1161 kg ha -1 . All the Ni application methods resulted in a 1.1-fold increase in the SPAD index, a 1.2-fold increase in photosynthesis, a 1.4-fold increase in nitrogenase, and a 3.9-fold increase in urease activity. Edaphoclimatic conditions exerted a significant influence on the treatments. The integrated approaches, namely, leaf application in conjunction with soil or seed fertilization, were more effective for enhancing yield in soybean cultivation systems. The determination of the ideal method is crucial for ensuring optimal absorption and utilization of this micronutrient and thus a feasible and sustainable management technology. Further research is warranted to establish official guidelines for the application of Ni in agricultural practices., (© 2024. The Author(s).)

The marine-based West Antarctic Ice Sheet (WAIS) is considered vulnerable to irreversible collapse under future climate trajectories, and its tipping point may lie within the mitigated warming scenarios of 1.5° to 2°C of the United Nations Paris Agreement. Knowledge of ice loss during similarly warm past climates could resolve this uncertainty, including the Last Interglacial when global sea levels were 5 to 10 meters higher than today and global average temperatures were 0.5° to 1.5°C warmer than preindustrial levels. Using a panel of genome-wide, single-nucleotide polymorphisms of a circum-Antarctic octopus, we show persistent, historic signals of gene flow only possible with complete WAIS collapse. Our results provide the first empirical evidence that the tipping point of WAIS loss could be reached even under stringent climate mitigation scenarios.

Studies of boron (B) and silicon (Si) synergy in cotton crops have shown promising results; however, the focus was on the foliar application of B and Si. Nonetheless, B is an element with little mobility in the plant and its best form of application is in the soil. Thus, the objective of this study was to evaluate the synergistic effect of soil applied B and foliar applied sSi on fiber quality and crop yield of cotton. For this purpose, a field experiment was carried out using cotton cultivar FM 985 GLTP. The soil's B in the experimental site is classified as low for cotton cultivation. The experiment was conducted in a randomized complete-block design, in a 3 × 2 factorial scheme, with three doses of B: 0.0 kg ha -1 (deficiency), 2.0 kg ha -1 (recommended dose), and 4.0 kg ha -1 (high dose) in the absence and presence (920 g L -1 ) of Si, with four replications. One week after the 4 th application of Si, B and Si leaf content was determined. At boll opening, crop yield was estimated, and fiber quality analysis was realized. Boron deficiency reduced cotton yield, in 11 and 9%, compared to the application of 2 and 4 kg ha -1 of B, respectively. The presence of Si, however, increased plant yield in 5% in the treatments with 0 and 2 kg ha -1 of B, respectively. Cotton fiber length and elongation were not influenced by the B doses and Si presence. Fiber breaking strength was increased in 5% by the presence of Si and was not influenced by B deficiency. Micronaire was 8% smaller in the treatment with 0 kg ha -1 of B and 6% smaller in the absence of Si. Short fiber index was 4% greater in the plants of the treatment with 0 kg ha -1 of B. The results of this study reports that the complementation with Si via foliar application increases fiber quality by enhance breaking strength and micronaire. In conclusion, the interaction between soil-applied B and foliar-applied Si is beneficial for cotton cultivation, resulting in high cotton yield with better fiber quality., (© 2023. The Author(s).)

In the present study, we aimed to produce CGP/PVA films containing entrapped anti-inflammatory drugs for wound dressing applications. Using a 3 3-1 fractional factorial design, the effect of each component was evaluated on the physicochemical and morphological properties of the produced materials. The best formulation for entrapment of diclofenac sodium and ketoprofen was also determined. The produced films presented high swelling capacity, with some formulations showing o porous structure. CGP/PVA films showed a maximum retention of 75.6% for diclofenac sodium and 32.2% for ketoprofen, and both drugs were released in a controlled manner for up to 7 h. The drug release kinetic was studied, and the data were fitted using a Korsmeyer-Peppas model, which suggested that the release mechanism is controlled by diffusion. These results indicate that CGP/PVA-based matrices have great potential to be used as drug-delivery systems for wound dressing applications, contributing to prolonging the drug's action time and then improving their anti-inflammatory efficacy., Competing Interests: Conflict of interestThe authors declare that they have no conflict of interest in the publication., (© King Abdulaziz City for Science and Technology 2023. Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.)

Understanding the drivers of evolutionary innovation provides a crucial perspective of how evolutionary processes unfold across taxa and ecological systems. It has been hypothesised that the Southern Ocean provided ecological opportunities for novelty in the past. However, the drivers of innovation are challenging to pinpoint as the evolutionary genetics of Southern Ocean fauna are influenced by Quaternary glacial-interglacial cycles, oceanic currents and species ecology. Here we examined the genome-wide single nucleotide polymorphisms of the Southern Ocean brittle stars Ophionotus victoriae (five arms, broadcaster) and O. hexactis (six arms, brooder). We found that O. victoriae and O. hexactis are closely-related species with interspecific gene flow. During the late Pleistocene, O. victoriae likely persisted in a connected deep water refugium and in situ refugia on the Antarctic continental shelf and around Antarctic islands; O. hexactis persisted exclusively within in situ island refugia. Within O. victoriae, contemporary gene flow linking to the Antarctic Circumpolar Current, regional gyres and other local oceanographic regimes was observed. Gene flow connecting West and East Antarctic islands near the Polar Front was also detected in O. hexactis. A strong association was detected between outlier loci and salinity in O. hexactis. Both O. victoriae and O. hexactis are associated with genome-wide increase in alleles at intermediate-frequencies; the alleles associated with this peak appear to be species specific, and these intermediate-frequency variants are far more excessive in O. hexactis. We hypothesise that the peak in alleles at intermediate frequencies could be related to adaptation in the recent past, linked to evolutionary innovations of increase in arm number and a switch to brooding from broadcasting, in O. hexactis., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

In this study, the antioxidant, antimicrobial, mechanical, optical, and barrier attributes of Solanum lycocarpum starch bio-based edible films incorporated with a phenolic extract from jaboticaba peel were investigated. Aiming to determine the effect of the polymers and the phenolic extract on the properties of the films, a three-factor simplex-lattice design was employed, and the formulation optimization was based on the produced films' antioxidant potential. The optimized formulation of the starch-PEJP film showed a reddish-pink color with no cracks or bubbles and 91% antioxidant activity against DPPH radical. The optimized starch-PEJP film showed good transparency properties and a potent UV-blocking action, presenting color variation as a function of the pH values. The optimized film was also considerably resistant and highly flexible, showing a water vapor permeability of 3.28 × 10 -6 g m -1 h -1 Pa -1 . The microbial permeation test and antimicrobial evaluation demonstrated that the optimized starch-PEJP film avoided microbial contamination and was potent in reducing the growth of Escherichia coli , Staphylococcus aureus , and Salmonella spp. In summary, the active starch-PEJP film showed great potential as an environmentally friendly and halochromic material, presenting antioxidant and antimicrobial properties and high UV-protecting activity.

The drivers behind evolutionary innovations such as contrasting life histories and morphological change are central questions of evolutionary biology. However, the environmental and ecological contexts linked to evolutionary innovations are generally unclear. During the Pleistocene glacial cycles, grounded ice sheets expanded across the Southern Ocean continental shelf. Limited ice-free areas remained, and fauna were isolated from other refugial populations. Survival in Southern Ocean refugia could present opportunities for ecological adaptation and evolutionary innovation. Here, we reconstructed the phylogeographic patterns of circum-Antarctic brittle stars Ophionotus victoriae and O .  hexactis with contrasting life histories (broadcasting vs brooding) and morphology (5 vs 6 arms). We examined the evolutionary relationship between the two species using cytochrome c oxidase subunit I (COI) data. COI data suggested that O .  victoriae is a single species (rather than a species complex) and is closely related to O .  hexactis (a separate species). Since their recent divergence in the mid-Pleistocene, O .  victoriae and O .  hexactis likely persisted differently throughout glacial maxima, in deep-sea and Antarctic island refugia, respectively. Genetic connectivity, within and between the Antarctic continental shelf and islands, was also observed and could be linked to the Antarctic Circumpolar Current and local oceanographic regimes. Signatures of a probable seascape corridor linking connectivity between the Scotia Sea and Prydz Bay are also highlighted. We suggest that survival in Antarctic island refugia was associated with increase in arm number and a switch from broadcast spawning to brooding in O .  hexactis , and propose that it could be linked to environmental changes (such as salinity) associated with intensified interglacial-glacial cycles., Competing Interests: The authors declare no competing interests., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The giant black tiger shrimp (Penaeus monodon) is native to the Indo-Pacific and is the second most farmed penaeid shrimp species globally. Understanding genetic structure, connectivity, and local adaptation among Indo-Pacific black tiger shrimp populations is important for informing sustainable fisheries management and aquaculture breeding programs. Population genetic and outlier detection analyses were undertaken using 10,593 genome-wide single nucleotide polymorphisms (SNPs) from 16 geographically disparate Indo-Pacific P. monodon populations. Levels of genetic diversity were highest for Southeast Asian populations and were lowest for Western Indian Ocean (WIO) populations. Both neutral (n = 9,930) and outlier (n = 663) loci datasets revealed a pattern of strong genetic structure of P. monodon corresponding with broad geographical regions and clear genetic breaks among samples within regions. Neutral loci revealed seven genetic clusters and the separation of Fiji and WIO clusters from all other clusters, whereas outlier loci revealed six genetic clusters and high genetic differentiation among populations. The neutral loci dataset estimated five migration events that indicated migration to Southeast Asia from the WIO, with partial connectivity to populations in both oceans. We also identified 26 putatively adaptive SNPs that exhibited significant Pearson correlation (P < 0.05) between minor allele frequency and maximum or minimum sea surface temperature. Matched transcriptome contig annotations suggest putatively adaptive SNPs involvement in cellular and metabolic processes, pigmentation, immune response, and currently unknown functions. This study provides novel genome-level insights that have direct implications for P. monodon aquaculture and fishery management practices., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

Nutritional deficiency is common in several regions of quinoa cultivation. Silicon (Si) can attenuate the stress caused by nutritional deficiency, but studies on the effects of Si supply on quinoa plants are still scarce. Given this scenario, our objective was to evaluate the symptoms in terms of tissue, physiological and nutritional effects of quinoa plants submitted to nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), and magnesium (Mg) deficiencies under Si presence. The experiment consisted of a factorial scheme 6 × 2, using a complete solution (CS), -N, -P, -K, -Ca, -Mg combined with absence and presence of Si (1.5 mmol L -1 ). Symptomatic, physiological, nutritional and evaluation vegetative were performed in quinoa crop. The deficiencies of N, P, K, Ca and Mg in quinoa cultivation caused visual symptoms characteristic of the deficiency caused by respective nutrients, hence decreasing the plant dry mass. However, Si supply attenuated the deficiency effects by preserving the photosynthetic apparatus, increasing the chlorophyll production, increasing the membrane integrity, and decreasing the electrolyte leakage. Thus, the Si supply attenuated the visual effects provided by deficiency of all nutrients, but stood out for N and Ca, because it reflected in a higher dry mass production. This occurred because, the Si promoted higher synthesis and protection of chlorophylls, and lower electrolyte leakage under Ca restriction, as well as decreased electrolyte leakage under N restriction., (© 2021. The Author(s).)

The New Zealand green-lipped mussel aquaculture industry is largely dependent on the supply of young mussels that wash up on Ninety Mile Beach (so-called Kaitaia spat), which are collected and trucked to aquaculture farms. The locations of source populations of Kaitaia spat are unknown and this lack of knowledge represents a major problem because spat supply may be irregular. We combined genotypic (microsatellite) and phenotypic (shell geochemistry) data in a geospatial framework to determine if this new approach can help identify source populations of mussels collected from two spat-collecting and four non-spat-collecting sites further south. Genetic analyses resolved differentiated clusters (mostly three clusters), but no obvious source populations. Shell geochemistry analyses resolved six differentiated clusters, as did the combined genotypic and phenotypic data. Analyses revealed high levels of spatial and temporal variability in the geochemistry signal. Whilst we have not been able to identify the source site(s) of Kaitaia spat our analyses indicate that geospatial testing using combined genotypic and phenotypic data is a powerful approach. Next steps should employ analyses of single nucleotide polymorphism markers with shell geochemistry and in conjunction with high resolution physical oceanographic modelling to resolve the longstanding question of the origin of Kaitaia spat.

Investigating historical gene flow in species complexes can indicate how environmental and reproductive barriers shape genome divergence during speciation. The processes influencing species diversification under environmental change remain one of the central focal points of evolutionary biology, particularly for marine organisms with high dispersal potential. We investigated genome-wide divergence, introgression patterns and inferred demographic history between species pairs of all six extant rock lobster species (Jasus spp.), which have a long larval duration of up to two years and have populated continental shelf and seamount habitats around the globe at approximately 40 o S. Genetic differentiation patterns reflected geographic isolation and the environment (i.e. habitat structure). Eastern Pacific species (J. caveorum and J. frontalis) were geographically more distant and genetically more differentiated from the remaining four species. Species associated with continental shelf habitats shared a common ancestry, but are geographically distant from one another. Similarly, species associated with island/seamount habitats in the Atlantic and Indian Oceans shared a common ancestry, but are also geographically distant. Benthic temperature was the environmental variable that explained most of the genetic differentiation (F ST ), while controlling for the effects of geographic distance. Eastern Pacific species retained a signal of strict isolation following ancient migration, whereas species pairs from Australia and Africa, and seamounts in the Indian and Atlantic oceans, included events of introgression after secondary contact. Our results reveal important effects of habitat and demographic processes on the recent divergence of species within the genus Jasus, providing one of the first empirical studies of genome-wide drivers of diversification that incorporates all extant species in a marine genus with long pelagic larval duration., (© 2020 John Wiley & Sons Ltd.)

While there is now strong evidence that many factors can shape dispersal, the mechanisms influencing connectivity patterns are species-specific and remain largely unknown for many species with a high dispersal potential. The rock lobsters Jasus tristani and Jasus paulensis have a long pelagic larval duration (up to 20 months) and inhabit seamounts and islands in the southern Atlantic and Indian Oceans, respectively. We used a multidisciplinary approach to assess the genetic relationships between J. tristani and J. paulensis , investigate historic and contemporary gene flow, and inform fisheries management. Using 17,256 neutral single nucleotide polymorphisms we found low but significant genetic differentiation. We show that patterns of connectivity changed over time in accordance with climatic fluctuations. Historic migration estimates showed stronger connectivity from the Indian to the Atlantic Ocean (influenced by the Agulhas Leakage). In contrast, the individual-based model coupled with contemporary migration estimates inferred from genetic data showed stronger inter-ocean connectivity in the opposite direction from the Atlantic to the Indian Ocean driven by the Subtropical Front. We suggest that the J. tristani and J. paulensis historical distribution might have extended further north (when water temperatures were lower) resulting in larval dispersal between the ocean basis being more influenced by the Agulhas Leakage than the Subtropical Front. As water temperatures in the region increase in accordance with anthropogenic climate change, a southern shift in the distribution range of J. tristani and J. paulensis could further reduce larval transport from the Indian to the Atlantic Ocean, adding complexity to fisheries management., Competing Interests: All authors declare that they have no competing interests., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Background: Restrictions to gene flow, genetic drift, and divergent selection associated with different environments are significant drivers of genetic differentiation. The black tiger shrimp (Penaeus monodon), is widely distributed throughout the Indian and Pacific Oceans including along the western, northern and eastern coastline of Australia, where it is an important aquaculture and fishery species. Understanding the genetic structure and the influence of environmental factors leading to adaptive differences among populations of this species is important for farm genetic improvement programs and sustainable fisheries management., Results: Based on 278 individuals obtained from seven geographically disparate Australian locations, 10,624 high-quality SNP loci were used to characterize genetic diversity, population structure, genetic connectivity, and adaptive divergence. Significant population structure and differentiation were revealed among wild populations (average F ST  = 0.001-0.107; p <  0.05). Eighty-nine putatively outlier SNPs were identified to be potentially associated with environmental variables by using both population differentiation (BayeScan and PCAdapt) and environmental association (redundancy analysis and latent factor mixed model) analysis methods. Clear population structure with similar spatial patterns were observed in both neutral and outlier markers with three genetically distinct groups identified (north Queensland, Northern Territory, and Western Australia). Redundancy, partial redundancy, and multiple regression on distance matrices analyses revealed that both geographical distance and environmental factors interact to generate the structure observed across Australian P. monodon populations., Conclusion: This study provides new insights on genetic population structure of Australian P. monodon in the face of environmental changes, which can be used to advance sustainable fisheries management and aquaculture breeding programs.

Photosynthetic efficiency has become the target of several breeding programs since the positive correlation between photosynthetic rate and yield in soybean suggests that the improvement of photosynthetic efficiency may be a promising target for new yield gains. However, studies on combining ability of soybean genotypes for physiological traits are still scarce in the literature. The objective of this study was to estimate the combining ability of soybean genotypes based on F2 generation aiming to identify superior parents and segregating populations for physiological traits. Twenty-eight F2 populations resulting from partial diallel crossings between eleven lines were evaluated in two crop seasons for the physiological traits: photosynthesis, stomatal conductance, internal CO2 concentration, and transpiration. General combining ability (GCA) of the parents and specific combining ability (SCA) of the F2 populations were estimated. Our findings reveal the predominance of additive effects in controlling the traits. The genotype TMG 7062 IPRO is the most promising parent for programs aiming at photosynthetic efficiency. We have also identified other promising parents and proposed cross-breeding with higher potential for obtaining superior lines for photosynthetic efficiency., Competing Interests: The authors have declared that no competing interests exist.

This study describes the use of X-ray fluorescence spectroscopy in Crotalaria ochroleuca seed technology. This work evaluated X-ray fluorescence techniques to estimate the physiological performance of different C. ochroleuca seed coat colours based on the concentration and distribution of Ca, P, K, and S in seed structures. The treatments consisted of seeds separated by coat colours (yellow, green, and red) and a control treatment (colour mix according to their natural occurrence in commercial lots), and was carried out in a completely randomized design, with four replications. The physiological performance was evaluated by analyzing the water content, germination, first germination count, germination speed index, electrical conductivity, seedling emergence, and seedling length and dry mass. X-ray fluorescence spectroscopy techniques were carried out with quantitative analyses (Ca, P, K, and S concentration in the seed coat and the whole seed) and qualitative analyses (macronutrient mapping). The EDXRF and μ-XRF techniques are efficient and promising to differentiate the physiological performance of C. ochroleuca seeds, based on the concentration and distribution of Ca, P, K, and S in different structures. Ca is predominant in the seed coat, and K, S, and P are found throughout the embryonic axis. Seeds of yellow and green coats have higher nutrients concentration and distribution in the embryonic axis, revealing high germinative capacity and physiological performance. Seeds of red coat have higher nutrients concentration in the seed coat and lower assimilation, showing less vigour, which interferes directly in the quality of commercial lots., Competing Interests: The authors have declared that no competing interests exist.

Background: The black-lip rock oyster (Saccostrea echinata) has considerable potential for aquaculture throughout the tropics. Previous attempts to farm S. echinata failed due to an insufficient supply of wild spat; however, the prospect of hatchery-based aquaculture has stimulated renewed interest, and small-scale farming is underway across northern Australia and in New Caledonia. The absence of knowledge surrounding the population genetic structure of this species has raised concerns about the genetic impacts of this emerging aquaculture industry. This study is the first to examine population genetics of S. echinata and employs both mitochondrial cytochrome c oxidase subunit I gene (COI) and single nucleotide polymorphism (SNP) markers., Results: The mitochondrial COI data set included 273 sequences of 594 base pair length, which comprised 74 haplotypes. The SNP data set included 27,887 filtered SNPs for 272 oysters and of these 31 SNPs were identified as candidate adaptive loci. Data from the mitochondrial COI analyses, supports a broad tropical Indo-Pacific distribution of S. echinata, and showed high haplotype and nucleotide diversities (0.887-1.000 and 0.005-0.008, respectively). Mitochondrial COI analyses also revealed a 'star-like' haplotype network, and significant and negative neutrality tests (Tajima's D = - 2.030, Fu's Fs = - 25.638, P < 0.001) support a recent population expansion after a bottleneck. The SNP analyses showed significant levels of population subdivision and four genetic clusters were identified: (1) the Noumea (New Caledonia) sample location; (2) the Bowen (north Queensland, Australia) sample location, and remaining sample locations in the Northern Territory, Australia (n = 8) were differentiated into two genetic clusters. These occurred at either side of the Wessel Islands and were termed (3) 'west' and (4) 'east' clusters, and two migrant individuals were detected between them. The SNP data showed a significant positive correlation between genetic and geographic distance (Mantel test, P < 0.001, R 2  = 0.798) and supported isolation by distance. Three candidate adaptive SNPs were identified as occurring within known genes and gene ontology was well described for the sex peptide receptor gene., Conclusions: Data supports the existence of genetically distinct populations of S. echinata, suggesting that management of wild and farmed stocks should be based upon multiple management units. This research has made information on population genetic structure and connectivity available for a new aquaculture species.

Knowledge about the underlying genetic architecture of phenotypic traits is needed to understand and predict evolutionary dynamics. The number of causal loci, magnitude of the effects and location in the genome are, however, still largely unknown. Here, we use genome-wide single-nucleotide polymorphism (SNP) data from two large-scale data sets on house sparrows and collared flycatchers to examine the genetic architecture of different morphological traits (tarsus length, wing length, body mass, bill depth, bill length, total and visible badge size and white wing patches). Genomic heritabilities were estimated using relatedness calculated from SNPs. The proportion of variance captured by the SNPs (SNP-based heritability) was lower in house sparrows compared with collared flycatchers, as expected given marker density (6348 SNPs in house sparrows versus 38 689 SNPs in collared flycatchers). Indeed, after downsampling to similar SNP density and sample size, this estimate was no longer markedly different between species. Chromosome-partitioning analyses demonstrated that the proportion of variance explained by each chromosome was significantly positively related to the chromosome size for some traits and, generally, that larger chromosomes tended to explain proportionally more variation than smaller chromosomes. Finally, we found two genome-wide significant associations with very small-effect sizes. One SNP on chromosome 20 was associated with bill length in house sparrows and explained 1.2% of phenotypic variation (V P ), and one SNP on chromosome 4 was associated with tarsus length in collared flycatchers (3% of V P ). Although we cannot exclude the possibility of undetected large-effect loci, our results indicate a polygenic basis for morphological traits.