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Persian walnut is a drought-sensitive species with considerable genetic variation in the photosynthesis and water use efficiency of its populations, which is largely unexplored. Here, we aimed to elucidate changes in the efficiency of photosynthesis and water content using a diverse panel of 60 walnut families which were submitted to a progressive drought for 24 days, followed by two weeks of re-watering. Severe water-withholding reduced leaf relative water content (RWC) by 20%, net photosynthetic rate (Pn) by 50%, stomatal conductance (gs) by 60%, intercellular CO2 concentration (Ci) by 30%, and transpiration rate (Tr) by 50%, but improved water use efficiency (WUE) by 25%. Severe water-withholding also inhibited photosystem II functionality as indicated by reduced quantum yield of intersystem electron transport (φEo) and transfer of electrons per reaction center (ET0/RC), also enhanced accumulation of QA (VJ) resulted in the reduction of the photosynthetic performance (PIABS) and maximal quantum yield of PSII (FV/FM); while elevated quantum yield of energy dissipation (φDo), energy fluxes for absorption (ABS/RC) and dissipated energy flux (DI0/RC) in walnut families. Cluster analysis classified families into three main groups (tolerant, moderately tolerant, and sensitive), with the tolerant group from dry climates exhibiting lesser alterations in assessed parameters than the other groups. Multivariate analysis of phenotypic data demonstrated that RWC and biophysical parameters related to the chlorophyll fluorescence such as FV/FM, φEo, φDo, PIABS, ABS/RC, ET0/RC, and DI0/RC represent fast, robust and non-destructive biomarkers for walnut performance under drought stress. Finally, phenotype-environment association analysis showed significant correlation of some photosynthetic traits with geoclimatic factors, suggesting a key role of climate and geography in the adaptation of walnut to its habitat conditions.

Uncovering the genetic basis of photosynthetic trait variation under drought stress is essential for breeding climate-resilient walnut cultivars. To this end, we examined photosynthetic capacity in a diverse panel of 150 walnut families (1500 seedlings) from various agro-climatic zones in their habitats and grown in a common garden experiment. Photosynthetic traits were measured under well-watered (WW), water-stressed (WS) and recovery (WR) conditions. We performed genome-wide association studies (GWAS) using three genomic datasets: genotyping by sequencing data (∼43 K SNPs) on both mother trees (MGBS) and progeny (PGBS) and the Axiom™ Juglans regia 700 K SNP array data (∼295 K SNPs) on mother trees (MArray). We identified 578 unique genomic regions linked with at least one trait in a specific treatment, 874 predicted genes that fell within 20 kb of a significant or suggestive SNP in at least two of the three GWAS datasets (MArray, MGBS, and PGBS), and 67 genes that fell within 20 kb of a significant SNP in all three GWAS datasets. Functional annotation identified several candidate pathways and genes that play crucial roles in photosynthesis, amino acid and carbohydrate metabolism, and signal transduction. Further network analysis identified 15 hub genes under WW, WS and WR conditions including GAPB, PSAN, CRR1, NTRC, DGD1, CYP38, and PETC which are involved in the photosynthetic responses. These findings shed light on possible strategies for improving walnut productivity under drought stress.

Walnut production is challenged by climate change and abiotic stresses. Elucidating the genomic basis of adaptation to climate is essential to breeding drought-tolerant cultivars for enhanced productivity in arid and semi-arid regions. Here, we aimed to identify loci potentially involved in water use efficiency (WUE) and adaptation to drought in Persian walnut using a diverse panel of 95 walnut families (950 seedlings) from Iran, which show contrasting levels of water availability in their native habitats. We analyzed associations between phenotypic, genotypic, and environmental variables from data sets of 609 000 high-quality single nucleotide polymorphisms (SNPs), three categories of phenotypic traits [WUE-related traits under drought, their drought stress index, and principal components (PCs)], and 21 climate variables and their combination (first three PCs). Our genotype–phenotype analysis identified 22 significant and 266 suggestive associations, some of which were for multiple traits, suggesting their correlation and a possible common genetic control. Also, genotype–environment association analysis found 115 significant and 265 suggestive SNP loci that displayed potential signals of local adaptation. Several sets of stress-responsive genes were found in the genomic regions significantly associated with the aforementioned traits. Most of the candidate genes identified are involved in abscisic acid signaling, stomatal regulation, transduction of environmental signals, antioxidant defense system, osmotic adjustment, and leaf growth and development. Upon validation, the marker–trait associations identified for drought tolerance-related traits would allow the selection and development of new walnut rootstocks or scion cultivars with superior WUE. [ABSTRACT FROM AUTHOR]

Simple sequence repeat (SSR) markers were used to authenticate ramets of 11 Persian walnut (Juglans regia L.) varieties. All varieties and 28 of their ramets (n = 39) were genotyped with 17 SSR markers. The genetic profiles revealed two off-types: the ramets Serr 4 (S4) and Vina 1 (V1). SSR fingerprints individuating 11 walnut varieties were possible using 13 polymorphic SSRs that could be used in the future to identify clones of these varieties. Except for ‘Chandler’, each cultivar could be distinguished using a combination of two SSR loci. This result emphasizes the efficacy of the SSR markers in true-to-type validation of walnut orchards. [ABSTRACT FROM AUTHOR]