Preharvest treatment with Bacillus velezensis HIII11 modulates cell wall dynamics and ripening-related processes in strawberry fruit.

Accurate genome assemblies are essential for comparative genomics, yet Hi-C-guided scaffolding can introduce structural errors that misrepresent chromosome architecture and bias evolutionary inferences. Here, we identified pervasive scaffolding errors - including artificial fusions, internal inversions, and incomplete contig mounting - in two previously published Lupinus genomes (L. cosentinii and L. digitatus) using a segmentation method based on LTR retrotransposon density. We re-assembled both genomes, producing chromosome-level references of 472.7 Mb (16 chromosomes) and 427.2 Mb (21 chromosomes), with BUSCO completeness >98.5%. Synteny validation and re-application of LTR profiling confirmed that all prior errors were resolved. Using these corrected genomes together with four additional Lupinus species and two outgroup legumes, we investigated post-polyploid evolution. Synonymous substitution rate (Ks) analysis revealed a genus-specific whole-genome duplication (WGD) event (Ks = 0.17) shared by all six Lupinus species. The proportion of WGD-derived genes varied markedly, from 60% in L. digitatus to only 36% in L. mutabilis, indicating differential diploidization. While all species retained a core set of WGD duplicates enriched in cytoskeleton organization, ion transport, and defense responses, each exhibited lineage-specific functional trajectories: cell wall modification in L. cosentinii and L. digitatus, nitrogen metabolism in L. albus and L. angustifolius, flower development in L. luteus, and stress/lipid metabolism in L. mutabilis. Our corrected assemblies provide optimal references for Lupinus comparative genomics, and our findings demonstrate that a shared WGD event can lead to both conserved and highly divergent post-polyploid fates, likely underpinning adaptive diversification within the genus.

Asymmetric cell division underpins cellular diversity in multicellular plants. These divisions are mechanosensitive, and preprophase band (PPB) formation hinges on cell-wall mechanical properties in plant cells. Yet, the spatial control mechanism governing this process in plants remains elusive. During grass stomatal development, mechanical cues originate from differential growth rates and cell wall modifications at the interface of guard mother cell/subsidiary mother cell (SMC). In this work, we have identified a maize receptor-like protein, KAI1, that functions as a master regulator of subsidiary cell formation within the stomatal complex. KAI1 governs division-plane orientation in SMCs through mechanochemical signaling: It perceives cell wall rigidity via pectin interaction, and subsequently recruits tubulin for PPB positioning, thereby directing division-plane specification. This work uncovers a plant-unique mechanosensitive protein that mediates extracellular matrix cues to cytoskeletal reorganization during asymmetric division for cell diversity. Our findings establish that KAI1 governs a cell wall mechanics-dependent PPB positioning to control the exact division plane alignment of the SMC. This mechanism subsequently mediates the regulation of SMC polarization and subsidiary cell morphogenesis during stomatal development.

Moso bamboo (Phyllostachys edulis) is a high-value renewable resource used for timber, fiber, and carbon sequestration, yet the cellular basis of its development remains poorly resolved. Here, we performed a systematic single-cell RNA-sequencing (scRNA-seq) analysis of moso bamboo seedlings, generating 31,658 high-quality single-cell transcriptomes. Initial unsupervised graph-based clustering partitioned the dataset into 23 clusters, which were subsequently assigned to 11 major cell-type categories through marker-guided and biologically informed annotation, including meristemoid, coleoptile L1 epidermis, cortex, epidermis, mesophyll precursor, mestome sheath, pericycle, phloem, endodermis, root cap, and xylem. Pseudo-time analysis suggested putative vascular-associated trajectories in which captured xylem and phloem cells were ordered along a bifurcating structure extending from meristem populations. Xylem cells were enriched in cell wall modification genes, while phloem cells preferentially expressed transport- and defense-related genes. Root cell differentiation exhibited polar patterning, progressing through seven stages from meristems to epidermis, endodermis, cortex, and root cap lineages. Leaf development followed a unidirectional trajectory from meristems to mestome sheath cells and coleoptile L1 epidermis, and ultimately to mesophyll cells. We further identified key regulatory factors underlying these cellular differentiations, revealing developmental path differences among roots, stems, and leaves. This study constructs a high-resolution cellular atlas of moso bamboo seedlings that delineates developmental programs underlying rapid growth and environmental adaptation, and uncovers dynamic trajectories from meristems to specialized cell types. These findings provide candidate regulators for future genetic studies to enhance biomass production and stress tolerance in moso bamboo.

Plants must constantly sense the status of their cell walls and respond by modifying cell walls to maintain growth and respond to developmental and external stimuli. This capacity to sense cell walls and mount appropriate responses are collectively called cell wall signaling. Reversible post-translational modifications, such as phosphorylation or ubiquitination, are often an important part of signal transduction since they can rapidly and reversibly modify protein function and/or localization. To uncover differential post-translational modifications during cell wall stress, we analyzed the ubiquitinated proteome of Arabidopsis seedlings treated with the cell wall synthesis inhibitor, isoxaben. We identified 49 proteins that are significantly differentially ubiquitinated under short-term isoxaben treatment. We examined the phenotypes of loss-of-function mutants affecting one of these candidates, UBQ6, and found that ubq6 mutants are relatively resistant to isoxaben. This resistance is partially conveyed by sustained cellulose synthesis during isoxaben treatment. We further found that UBQ6 is a ubiquitin extension protein family member and that it is cleaved into two domains: the ubiquitin-like domain and the ribosomal-like domain. We confirmed that the site that is differentially ubiquitinated under cell wall stress is within the ribosomal-like domain of UBQ6 and found that the ribosomal-like domain is required for wild-type cell wall stress responses. We propose a model in which ubiquitination of the ribosomal-like domain of a UBQ6 has a specific role in cell wall signaling and cell wall stress responses, independent of the ubiquitin molecule created by UBQ6 protein cleavage.

Root hairs are specialized extensions of root epidermal cells that allow plants to explore and attach to the soil. They exhibit polar growth under the influence of isotropic turgor pressure thanks to the anisotropic nature of their cell walls. This unidirectional growth is regulated by myriad subcellular factors such as microtubule and actin dynamics, a tip-focused calcium gradient, and the interplays between gradients of apoplastic and cytosolic pH and reactive oxygen species. All these players also influence cell wall dynamics by forming feedback loops that modulate cell wall assembly and modification, which are essential processes for root hair morphogenesis. In this review we discuss the functions of cell wall polysaccharides and proteins and their impacts on the biomechanics of root hair growth at each developmental stage. We also discuss important open questions and technical advancements in studying root hair mechanobiology. Despite significant progress, many of the spatiotemporal changes that occur in the cell walls of root hairs remain undiscovered. Therefore, we highlight ongoing research and exciting future avenues that will shed light on cell wall dynamics, biomechanics, and mechanobiology of root hair morphogenesis.

Black rot, caused by Phytophthora parasitica, poses a significant threat to the commercial cultivation of Dendrobium spp., particularly the cultivar 'Earsakul'. However, the black rot resistance mechanisms in Dendrobium Sonia 'Earsakul' remain unexplored. This study elucidates the molecular mechanisms underlying black rot resistance by comparing a resistant ethyl methanesulfonate (EMS)-mutagenized line, SUT13E18301 with a susceptible non-mutagenized control, SUT16C014 in control (without pathogen inoculation) and inoculated conditions. Resistance phenotyping using detached leaf assay revealed stark contrasts in disease severity, prompting comprehensive transcriptomic analyses via RNA sequencing at 12- and 24-hours post-inoculation (hpi), compared with control condition. Bioinformatic analysis identified 4,190 significantly differentially expressed genes (DEGs), most of which were associated with defense responses. At the early stage (12 hpi), the genes related to signaling pathway, transcriptional reprogramming, and cell wall modification were differentially expressed. At the later infection stage (24 hpi), results highlighted genes with important roles in pathogen recognition, hormone signaling, cell wall modification, antimicrobial compounds/defense proteins production, hypersensitive response, and detoxification, all related to disease resistance mechanisms. Among these, nearly all of twenty-three candidate genes, including peroxidase (POD) 51-like, beta-glucosidase 11-like, pectinesterase, chitinase 2-like, transcription factor MYB6, and fasciclin-like arabinogalactan protein 11 (FLA11) showed up-regulation in the resistant line, whereas reduced responses in the susceptible line. Quantitative real-time PCR (qPCR) validation confirmed these expression patterns, showing strong positive correlation (R = 0.94). Furthermore, a highly significant increase in POD activity was observed at 24 hpi compared to 0 and 12 hpi in the resistant line. Conversely, the susceptible line showed no significant differences across the time points. Additionally, the weighted gene co-expression network analysis (WGCNA) and the regulatory network of candidate genes indicated that resistance is mediated by complex interactions among multiple regulatory components, rather than a single predominant pathway, and integration of structural and chemical defenses. These findings provide crucial insights into the genetic basis of black rot resistance and identify valuable molecular targets for breeding black rot resistance inDendrobium.

Corolla abscission is an important agronomic trait that affects fruit quality and disease susceptibility in blueberry. However, the underlying molecular mechanisms remain poorly understood. This study aimed to investigate the molecular and physiological factors associated with corolla abscission by comparing two blueberry cultivars with contrasting corolla abscission characteristics: the caducous 'Magica' ('F6') and the persistent 'Eureka-Sunrise' ('L25'). Cytological analysis revealed distinct structural differences in the abscission zone (AZ) between the two cultivars. In 'F6', cell separation occurred after full bloom, whereas AZ cells in 'L25' remained intact and gradually resembled adjacent tissues. To explore the molecular basis of these differences, transcriptome analysis was performed at three developmental stages. Differential expression and enrichment analyses indicated that genes related to plant hormone signaling, cell wall organization, and oxidative stress response were consistently involved. Weighted gene co-expression network analysis further identified modules associated with cultivar-specific traits, with genes related to ethylene and abscisic acid pathways showing higher expression in 'F6', while those associated with auxin metabolism and stress-related processes were more active in 'L25'. To validate these findings, the activities of cell wall-related enzymes were measured. Polygalacturonase activity increased at the stage corresponding to abscission initiation in 'F6', while remaining relatively low in 'L25'. In addition, cellulase activity was consistently higher in 'F6' than in 'L25'. These results indicate that corolla abscission in blueberry is associated with coordinated changes in hormone-related pathways and cell wall modification. Differences in the regulation of these processes between cultivars may contribute to variation in abscission behavior.

Mineral (micro- and macronutrient)-doped quantum fertilizers developed via green synthesis represent a novel and transformative paradigm in sustainable agriculture, offering an ecofriendly strategy to maximize nutrient use efficiency, stimulate plant growth, and significantly reduce dependence on conventional chemical fertilizers. Here, we report the synthesis of Mg-, Zn-, Fe- and B-doped carbon quantum dots via a microwave-assisted method using oxalic acid as a molecular precursor. Notably, boron-doped carbon quantum dots (B-QDs) exhibit dual roles of plant growth promotion and significant antibacterial activity, as comprehensively characterized and functionally validated. Structural analyses (UV-vis spectroscopy, TEM, XPS, and fluorescence spectroscopy) confirmed the formation of quantum dots with an average diameter of 8–12 nm. B-QD treatments significantly enhanced seed germination, root elongation, and root number compared with the control (without mineral), boron alone, and Mg-doped and Fe-doped carbon quantum dots (Mg-QDs and Fe-QDs, respectively). However, Zn-doped carbon quantum dots (Zn-QDs) showed better seed germination and root elongation ability, but no antibacterial activity was observed. Fluorescence imaging further demonstrated efficient internalization and distribution of B-QDs in root tissues, while binding studies revealed strong interactions with pectic acid, a major polysaccharide in the cell walls suggesting a role in modulating cell wall dynamics. Antibacterial assays confirmed selective inhibition of phytopathogens such as Pseudomonas syringae and Ralstonia solanacearum without adversely affecting beneficial soil microbes. Furthermore, B-QDs displayed potent antibiofilm activity, validating their dual function as both plant growth promoters and eco-friendly antimicrobial agents. These seminal findings catalyze a new scientific quest to harness the power of “quantum fertilizers” toward advanced sustainable agriculture and next-generation plant disease management.

Dataset of the changes in sugarbeet taproot cell wall composition, firmness, and cell wall-related gene expression as a function of postharvest storage time and temperature.

Calcium attenuates cadmium toxicity in Cosmos bipinnatus through cell wall remodeling and metabolic regulation: A transcriptomic and physiological analysis.

Acidic soils trigger a surge of soluble aluminum (Al), thereby causing Al3+ toxicity that inhibits plant growth and limits crop yields. Rhodomyrtus tomentosa, a shrub distributed in tropical and subtropical regions with acid soil, is an important genetic source for exploring the mechanism of Al tolerance. The xyloglucan endotransglucosylase/hydrolase (XTH) cleaves and rejoins xyloglucan polymers, which is important for plants to adapt to Al3+ stress. In the present study, a total of 29 RtXTHs were identified in R. tomentosa. These RtXTHs showed distinct tissue-specific expression patterns. Al3+ stress significantly upregulated several RtXTH genes in the roots, among which RtXTH2 showed the most prominent responsiveness. The subcellular localization assay determined that RtXTH2 encoded a cell wall-localized XTH protein. Overexpression of RtXTH2 enhanced Al tolerance in Arabidopsis by reducing hemicellulose and pectin contents in the cell wall. RtERF1, an Al3+-induced AP2/ERF transcription factor, directly activated RtXTH2 by binding to the CRT elements in its promoter. Silencing either RtXTH2 or RtERF1 reduced Al tolerance in R. tomentosa seedlings, as evidenced by increased Al accumulation, enhanced membrane damage, and more severe inhibition of root elongation under Al3+ stress. In summary, our results indicate that RtERF1 activates RtXTH2 to modify cell wall composition, thereby enhancing Al tolerance in R. tomentosa.

Boron (B) is an essential micronutrient critical for plant growth and reproductive development, primarily through its role in the cell wall. Calcium (Ca2+) similarly stabilizes cell wall architecture by forming cross-links with de-esterified pectin. In Arabidopsis thaliana, B deficiency rapidly induces JASMO0NATE RESISTANT 1 (JAR1) and jasmonic acid (JA) accumulation, which negatively regulates growth. However, whether and how JAR1-mediated JA signaling modulates cell wall B and Ca2+ partitioning to confer B-deficiency tolerance remains unclear. Here, we characterized the function of BnaA5.JAR1 in rapeseed (Brassica napus L.) under B deficiency. Pharmacological inhibition of JA biosynthesis with ibuprofen partially alleviated B-deficiency-induced shoot growth inhibition. BnaA5.JAR1 transcript levels were rapidly and strongly induced by B deprivation. CRISPR/Cas9-mediated knockout of BnaA5.JAR1 enhanced tolerance to B deficiency, whereas overexpression increased sensitivity, despite unchanged leaf B concentrations. Notably, mutant lines maintained robust tolerance throughout the reproductive stage, effectively rescuing the 'flowering without seed setting' phenotype characteristic of B deficiency. Under B limitation, these knockout lines retained higher B and Ca2+ concentrations in alkali-soluble pectin, exhibited a lower degree of pectin methylesterification, and maintained thinner, more structurally normal cell walls compared with overexpression lines. Exogenous Ca2+ mitigated B deficiency symptoms without increasing leaf B concentration. Collectively, these findings establish BnaA5.JAR1 as a negative regulator of B-deficiency tolerance and demonstrate that suppressing its activity enhances Ca2+-mediated cell wall stabilization across vegetative and reproductive stages, offering a targeted molecular strategy for breeding B-efficient rapeseed cultivars.

Although lychee peel extract (LPE) is rich in bioactive compounds, its potential for postharvest fruit preservation remains unexplored. We hypothesised that LPE would act synergistically with chitosan (CH) to delay mango ripening by simultaneously modulating cell wall integrity, pigment metabolism, and hormone signaling pathways. Here, we demonstrate that chitosan combined with lychee peel extract (CHL) delays mango ripening through a multi-targeted mechanism. Specifically, CHL outperformed chitosan alone by significantly suppressing peel yellowing, maintaining fruit firmness, and reducing decay over 12days of storage. Integrated transcriptomic and metabolomic analyses revealed that LPE reprogrammed ripening-associated pathways by (1) upregulating cell wall remodeling genes (CSLE1, XTH23) to stabilize pectin architecture, (2) retaining chlorophyll via suppressed CRTISO and PSY (carotenoid synthesis) and enhanced CHLP (chlorophyll biosynthesis), and (3) decoupling sugar-acid dynamics through γ-aminobutyric acid (GABA) and succinic acid accumulation. Notably, LPE attenuated ethylene-auxin- abscisic acid (ABA) crosstalk by downregulating ripening-specific transcription factors (ERF003, bZIPs) while activating stress-responsive WRKYs. These findings establish LPE as a sustainable alternative to synthetic preservatives, leveraging agricultural byproducts for eco-friendly fruit preservation.

Molybdenum-selenium doped carbon dots enhance cadmium detoxification in lettuce through root remodeling and multi-pathway regulation.

Tuber eye depth is a key agronomic trait in potato, influencing both processing efficiency and market appearance. To investigate the genetic basis of deep-eye phenotype in the elite Chinese tetraploid cultivar Jinshu 16, we constructed an F1 segregating population by crossing Jinshu 16 (deep eye) with Atlantic (shallow eye). Phenotypic evaluation across two seasons revealed a 1:1 segregation ratio for eye depth (Eyd), indicative of control by a single dominant locus. By integrating whole-genome resequencing and bulk segregant analysis (BSA-seq) of extreme phenotypes, we fine-mapped the major locus to a 49.0-50.7Mb interval on chromosome 10, co-localizing with the previously reported Eyd locus. Cytological examination showed increased cell density in the eye region of Jinshu 16 tubers. Transcriptome analysis identified specific upregulation of several lipid transfer protein and peroxidase genes within the mapped region in Jinshu 16 eye tissues, implicating lipid metabolism and cell wall modification in eye depth determination. Developed Insertion‑Deletion (InDel) markers from this region were significantly associated with the phenotype in the population. Our study defines the genetic locus responsible for the deep-eye trait in Jinshu 16, providing a foundation for gene functional studies and molecular marker-assisted breeding in this cultivar.

Dwarfism is crucial for intensive cultivation and labor-saving management in modern orchards. However, the molecular mechanisms underlying pear tree dwarfing remain largely unclear. Here, we performed comparative transcriptome analysis between dwarf pear germplasm and pear cultivar 'Cuiguan'. The dwarf pear germplasm exhibited significantly shorter internode and branch length compared to 'Cuiguan'. Histological analysis revealed that cortical cells in the dwarf pear germplasm were disordered and irregular, and longer than those of 'Cuiguan'. Comparative transcriptome analysis of shoot apex from young shoots of the dwarf germplasm and 'Cuiguan' was conducted and a total of 13,169 differentially expressed genes (DEGs) were identified. Functional enrichment analysis revealed that functional terms related to plant hormone biosynthesis and signal transduction were overrepresented in DEGs. Genes involved in brassinosteroid (BR) and gibberellin (GA) biosynthesis were downregulated in the dwarf germplasm. DEGs involved in transcription regulation and cell wall formation were also identified, which play potential roles in tree dwarfing. In addition, expression level of genes within the chromosomal region containing PcDw locus, which has been reported as the dominant gene controlling dwarf trait, was investigated in both the dwarf pear germplasm and 'Cuiguan'. Based on comparative analysis, nine genes in this region are considered to be closely associated with dwarf traits. The dwarf germplasm and 'Cuiguan' showed distinct tissue structure, with dwarfism mainly associated with defects in cell division and elongation. Comparative transcriptome analysis identified a set of DEGs involved in plant hormones biosynthesis, signaling and transport, as well as cell wall biosynthesis, modification and degradation. By integrating transcriptome data with information related to the PcDW locus, nine genes were identified as key candidates. Overall, the results of this study provide insights for understanding the mechanisms underlying pear dwarfism.

Anthracnose disease, caused by Colletotrichum species, poses a significant threat to global litchi production, yet the molecular mechanisms governing host resistance remain poorly understood. To dissect the genetic basis of anthracnose resistance, we employed a comparative transcriptomic approach using two contrasting cultivars: 'YuJinQiu' (DR, disease-resistant genotype) and 'BaiTangYing' (DS, disease-susceptible genotype). Field evaluations and controlled infection assays demonstrated evident phenotypic divergence, with DR exhibiting delayed disease progression and 43.7% smaller lesion areas compared to DS at 72h post-inoculation (hpi). Time-resolved RNA sequencing (0-72 hpi) revealed genotype-specific transcriptional dynamics, where DR displayed fewer differentially expressed genes (DEGs; 819-1457) compared to DS (5195-5735), suggesting a more targeted and efficient defense response. Functional enrichment analyses highlighted rapid activation of innate immunity pathways in DR, including pattern-triggered immunity, MAPK signaling, and jasmonic acid/ethylene biosynthesis, whereas DS prioritized cell wall modification and compensatory secondary metabolic processes. Weighted gene co-expression network analysis (WGCNA) pinpointed three modules tightly linked to anthracnose resistance, enriched for receptor-like kinases (RLKs), nucleotide-binding leucine-rich repeat (NLR) proteins, and phenylpropanoid biosynthesis genes. Hub regulators, including WRKY transcription factors (e.g., WRKY33), ubiquitin ligases, and pathogenesis-related proteins (PR1, PR5), were identified as central coordinators of defense signaling. Strikingly, DR exhibited sustained upregulation of effector-triggered immunity markers, particularly nucleotide-binding leucine-rich repeat (NLR) genes, and early accumulation of phytoalexins, correlating with pathogen suppression. Experimental validation via qRT-PCR confirmed the reliability of transcriptomic data. Our study unravels the multilayer regulatory network underlying litchi anthracnose resistance, providing not only a mechanistic model of cultivar-specific responses but also a robust gene toolkit for accelerating the development of resistant cultivars through marker-assisted breeding.

Grain length is a crucial determinant of rice yield and quality. Although von Willebrand factor type A (VWA) domain proteins have recently emerged as regulators of plant architecture in cereals, their roles in controlling rice grain length remain largely unexplored. Here, we report the functional characterization of OsvWA36, a VWA-domain protein selected from a TMT-based quantitative proteomic screen of indica rice varieties differing in grain length. Loss-of-function OsvWA36 mutants exhibited significantly reduced grain length and thousand-grain weight, while complementation and overexpression assays confirmed its positive regulatory role in these traits. Cytological analysis revealed that the shortened grain phenotype was due to suppressed longitudinal elongation of hull epidermal cells, which was accompanied by aberrantly enhanced lignin deposition. Transcriptomic profiling and Gene Ontology (GO) enrichment analysis demonstrated that OsvWA36 is essential for the expression of a comprehensive suite of cell wall biosynthesis and modification genes, including those involved in cellulose, pectin, and lignin metabolism. Furthermore, OsvWA36 localizes to punctate structures on the endoplasmic reticulum (ER). Our study establishes OsvWA36 as a novel VWA-domain protein that positively regulates grain length by orchestrating cell wall remodeling programs, thereby bridging VWA protein function with the transcriptional regulation of cell wall dynamics in rice. This work not only identifies a promising genetic target for molecular breeding but also provides a new molecular framework for understanding grain size regulation in cereal crops.

HaEXLA2 and HaEXLB2 enhance drought tolerance but suppress lignin defense, increasing broomrape susceptibility, revealing a trade-off in sunflowers. Expansins modulate cell wall dynamics to mediate plant stress responses. This study presents the first genome-wide analysis of the expansin gene-family in the sunflower (Helianthus annuus) and sunflower broomrape (Orobanche cumana) pathosystem, identifying 51HaEXPand 23OcEXPgenes. Phylogenetic classification placed them into four canonical subfamilies: expansin A (EXPA), expansin B (EXPB), expansin-like A (EXLA), and expansin-like B (EXLB). The family's expansion was primarily driven by tandem duplication events under strong purifying selection. Promoter cis-element analysis revealed a high abundance of stress-responsive elements, predicting roles in ABA, JA, and hypoxia signaling. RNA-seq and RT-qPCR validation demonstrated that HaEXLA2andHaEXLB2were distinctly upregulated in resistant sunflowers during earlyO. cumanaparasitization. Contrary to expectations, functional characterization revealed that their transient overexpressionsuppressed the phenylpropanoid pathway, downregulating key lignin biosynthetic genes (PAL1, 4CL2, COMT, and CAD1) and reducing lignin accumulation. This suppression of lignin-based defensesignificantly increased the susceptibilityof otherwise resistant sunflower cultivars to broomrape. In parallel, heterologous expression ofHaEXLA2andHaEXLB2in yeast enhanced drought tolerance by modulating cell wall properties. Our findings reveal a critical trade-off:HaEXLA2andHaEXLB2enhance abiotic stress adaptation while simultaneously suppressing lignin-based biotic defense against a root parasite. This dual functionality suggests them as potential targets for engineering stress resilience, though their negative impact on parasite resistance must be addressed in applied breeding programs.