\n Dioecious plants exhibit sexual dimorphism in both sexual features (reproductive organs) and secondary sex characteristics (vegetative traits). Sexual differences in secondary traits, including morphological, physiological and ecological characters, have been commonly associated with trade-offs between the cost of reproduction and other plant functions. Such trade-offs may be modified by environmental stressors, although there is evidence that sexually dimorphic responses to stress do not always exist in all plant species. When sexual dimorphism exists, sexually different responses appear to depend on the species and stress types. Yet, further studies on dioecious plant species are needed to allow the generalization of stress effects on males and females. Additionally, sexual dimorphism may influence the frequency and distribution of the sexes along environmental gradients, likely causing niche differentiation and spatial segregation of sexes. At the present, the causes and mechanisms governing sex ratio biases are poorly understood. This review aims to discuss sex-specific responses and sex ratio biases occurring under adverse conditions, which will advance our knowledge of sexually dimorphic responses to environmental stressors.

Sexual differences and sex ratios of dioecious plants under stressful environments

Effects of climate warming on tree growth and physiology may be driven by direct thermal effects and/or by changes in soil moisture. Dioecious tree species usually show sexual spatial segregation along abiotic gradients; however, few studies have assessed the sex-specific responses to warming in dioecious trees. We investigated the sex-specific responses in growth, photosynthesis, nonstructural carbohydrate (NSC), water-use efficiency and whole-plant hydraulic conductance (KP) of the dioecious tree species Populus cathayana Rehd. under +4 °C elevated temperature with and without supplemental water. For both sexes, high-temperature treatments significantly decreased growth (height and biomass), photosynthetic rate (A), the ratio of A to dark respiration rate, stomatal conductance (gs), transpiration rate, NSC, leaf water potential and KP, but increased water-use efficiency (estimated from carbon isotope composition). Under warming with supplemental water, most traits of females did not change relative to ambient conditions, but traits of males decreased, resulting in greater sexual differences. Females showed a lower KP, and their gs and A responded more steeply with water-related traits than males. These results show that the effect of summer warming on growth and photosynthesis was driven mainly by soil moisture in female P. cathayana, while male performance was mainly related to temperature. Females may experience less thermal stress than males due to flexible water balance strategy via stomata regulation and water use.

Effect of summer warming on growth, photosynthesis and water status in female and male Populus cathayana: implications for sex-specific drought and heat tolerances.

Sexual dimorphism occurs regarding carbon and nitrogen metabolic processes in response to nitrogen supply. Differences in fixation and remobilization of carbon and allocation and assimilation of nitrogen between sexes may differ under severe defoliation. The dioecious species Populus cathayana was studied after two defoliation treatments with two N levels. Males had a higher capacity of carbon fixation because of higher gas exchange and fluorescence traits of leaves after severe long-term defoliation under deficient N. Males had higher leaf abscisic acid, stomatal conductance and leaf sucrose phosphate synthase activity increasing transport of sucrose to sinks. Males had a higher carbon sink than females, because under N-deficient conditions, males accumulated >131.10% and 90.65% root starch than males in the control, whereas females accumulated >40.55% and 52.81%, respectively, than females in the control group. Males allocated less non-protein N (NNon-p) to leaves, having higher nitrogen use efficiency (photosynthetic nitrogen use efficiency), higher glutamate dehydrogenase (GDH) and higher leaf GDH expression, even after long-term severe defoliation under deficient N. Females had higher leaf jasmonic acid concentration and NNon-p. The present study suggested that females allocated more carbon and nitrogen to defense chemicals than males after long-term severe defoliation under deficient N.

Differences in carbon and nitrogen metabolism between male and female Populus cathayana in response to deficient nitrogen

Different sexual impacts of dioecious Populus euphratica on microbial communities and nitrogen cycle processes in natural forests

Dioecious plants have coevolved with diverse plant microbiomes, which are crucial for the fitness and productivity of their host. Sexual dimorphism in morphology, physiology, or gene expression may relate to different microbial compositions that affect male and female fitness in different environments. However, sex-specific impacts on ecological processes that control the microbiome assembly are not well known. In this study, Populus cathayana males and females were planted in different nitrogen conditions. It was hypothesized that males and females differently affect bacterial and fungal communities in the rhizosphere soil, roots, old leaves, and young leaves. Physiological traits and transcriptome profiles of male and female plants were investigated to reveal potential mechanisms that control the microbiome assembly.Our results showed strong niche differentiation that shapes microbial communities leading to a rapid loss of diversity along a decreasing pH gradient from the rhizosphere soil to leaves. Sex had different impacts on the microbial assembly in each niche. Especially fungal endophytes showed great differences in the community structure, keystone species, and community complexity between P. cathayana males and females. For example, the fungal co-occurrence network was more complex and the alpha diversity was significantly higher in young female leaves compared to young male leaves. Transcriptome profiles revealed substantial differences in plant-pathogen interactions and physiological traits that clearly demonstrated divergent internal environments for endophytes inhabiting males and females. Starch and pH of young leaves significantly affected the abundance of Proteobacteria, while tannin and pH of roots showed significant effects on the abundance of Chloroflexi, Actinobacteria, and Proteobacteria, and on the bacterial Shannon diversity.Our results provided important knowledge for understanding sexual dimorphism that affects microbial assemblies, thus advancing our understanding of plant-microbiome interactions. Video Abstract. 

/pdf/plant-sex-affects-plant-microbiome-assemblies-of-dioecious-2mrwhwr0.pdf

Plant sex affects plant-microbiome assemblies of dioecious Populus cathayana trees under different soil nitrogen conditions

Soil phosphorus (P) availability and its distribution influence plant growth and productivity, but how they affect the growth dynamics and sex-specific P acquisition strategies of dioecious plant species is poorly understood. In this study, the impact of soil P availability and its distribution on dioecious Populus cathayana was characterized. P. cathayana males and females were grown under three levels of P supply, and with homogeneous or heterogeneous P distribution. Females had a greater total root length, specific root length (SRL), biomass and foliar P concentration under high P supply. Under P deficiency, males had a smaller root system than females but a greater exudation of soil acid phosphatase, and a higher colonization rate and arbuscular mycorrhizal hyphal biomass, suggesting a better capacity to mine P and a stronger association with arbuscular mycorrhizal fungi to forage P. Heterogeneous P distribution enhanced growth and root length density (RLD) in females. Female root proliferation in P-rich patches was related to increased foliar P assimilation. Localized P application for increasing P availability did not enhance the biomass accumulation and the morphological plasticity of roots in males, but it raised hyphal biomass. The findings herein indicate that sex-specific strategies in P acquisition relate to root morphology, root exudation and mycorrhizal symbioses, and they may contribute to sex-specific resource utilization patterns and niche segregation.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.16170

Sex-specific strategies of phosphorus (P) acquisition in Populus cathayana as affected by soil P availability and distribution

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Sex-specific strategies of phosphorus (P) acquisition in Populus cathayana as affected by soil P availability and distribution