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- Brachypodium hybridum (2)
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- Brachypodium distachyon (1)
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Long and short-term climatic variation affect the ability of plants to simultaneously cope with increasing abiotic stress and biotic interactions. Specifically, ecotypes adapted to different climatic conditions (i.e., long-term legacy) may have to adjust their allocation to chemical defenses against enemies under acute drought (i.e., short-term response). Although several studies have addressed drought effects on chemical defense production, little is known about their intraspecific variation along resource gradients. Studying intraspecific variation is important for understanding how different environments select for defense strategies and how these may be affected directly and indirectly by changing climatic conditions. We conducted greenhouse experiments with the annual Biscutella didyma (Brassicaceae) to test the effects of long-term climatic legacy versus short-term drought stress on the concentrations of defense compounds (glucosinolates). To this aim, four ecotypes originating from a steep aridity gradient were exposed to contrasting water treatments. Concentrations of chemical defenses were measured separately in leaves of young (8 weeks) and old (14 weeks) plants, respectively. For young plants, ecotypes from the wettest climate (long-term legacy) as well as plants receiving high water treatments (short-term response) were better defended. A marginally significant interaction suggested that wetter ecotypes experienced a larger shift in defense production across water treatments. Older plants contained much lower glucosinolate concentrations and showed no differences between ecotypes and water treatments. Our results indicate that younger plants invest more resources into chemical defenses, possibly due to higher vulnerability to tissue loss compared to older plants. We propose that the strong response of wet ecotypes to water availability may be explained by a less pronounced adaptation to drought.
Clinal population divergence in an adaptive parental environmental effect that adjusts seed banking
(2017)
Bet-hedging via between-year seed dormancy is a costly strategy for plants in unpredictable environments. Theoretically, fitness costs can be reduced through a parental environmental effect when the environment is partly predictable. We tested whether populations from environments that differ in predictability diverged in parental effects on seed dormancy. Common garden-produced seeds of the two annual plant species Biscutella didyma and Bromus fasciculatus collected along an aridity gradient were grown under 12 irrigation treatments. Offspring germination was evaluated and related to environmental correlations between generations and their fitness consequences at the four study sites. One species exhibited strong seed dormancy that increased with unpredictability in seasonal precipitation. The parental effect on seed dormancy also increased proportionally with the environmental correlation between precipitation in the parental season and seedling density in the following season; this correlation increased from mesic to arid environments. Because fitness was negatively related to density, this parental effect may be adaptive. However, the lack of dormancy in the second species indicates that bet-hedging is not the only strategy for annual plants in arid environments. Our results provide the first evidence for clinal variation in the relative strength of parental effects along environmental gradients.
This study tested systematically at two spatial scales for key traits shaping within-species ecotypic differentiation under increasing aridity. It assessed different plant strategy theories and considered potential implications for climate change. We studied the widespread Mediterranean grass Brachypodium hybridum. At large scale, we tested 14 populations along a steep natural aridity gradient (114-954 mm annual rainfall). At small scale, we tested the microclimatic contrast between plants originating from corresponding north (more mesic) and south (more arid) exposed hillslopes. Fifteen traits were measured in the greenhouse, including the popular traits of the LeafHeight- Seed scheme (SLA, plant height, seed mass), several traits on phenology, architecture, growth, fitness, and rarely measured root traits. Clear trait shifts indicated ecotypic differentiation along the large-scale gradient. Earlier phenology, higher reproductive allocation and reduced root investment characterized arid ecotypes. Surprisingly, no trait of the Leaf-Height-Seed scheme shifted with aridity and root responses were opposite to the theory of optimal resource partitioning. Trait differences between north and south exposures were small, often inconsistent between sites, and poorly matched the trends across the large-scale gradient. South exposures thus appeared unlikely to harbour distinct ecotypes better adapted to aridity. Our findings highlight ecotypes as a crucial way how species span environmental gradients, yet underpinning their restriction at small spatial scales. In combination, this possibly renders populations more vulnerable to climate change. We draw attention to specific, partly unexpected traits and pose the question whether the LeafHeight- Seed scheme has limited applicability for intraspecific investigations in drylands.
The phylogenetic structure of communities (PSC) reveals how evolutionary history affects community assembly processes. However, there are important knowledge gaps on PSC patterns for annual communities and there is a need for studies along environmental gradients in dry ecosystems where several processes shape PSC. Here, we investigated the PSC of annual plants along an aridity gradient in Israel, including eight years, two spatial scales, the effects of shrubs on understory, and the phylogenetic signal of important traits. Increasing drought stress led to overdispersed PSC at the drier end of the gradient, indicating that species were less related than expected by chance. This was supported at a smaller spatial scale, where within the drier sites, communities in open- more arid- habitats were more overdispersed than those under nurse shrubs. Interestingly, some key traits related to drought resistance were not conserved in the phylogeny. Together, our findings suggested that while habitat filtering selected for drought resistance strategies, these strategies evolved independently along multiple contrasting evolutionary lineages. Our comprehensive PSC study provides strong evidence for the interacting effects of habitat filtering and plant- plant interactions, particularly highlighting that the conservative evolution of traits should not be assumed in future interpretations of PSC patterns.
The ecological benefits of polyploidy are intensely debated. Some authors argue that plants with duplicated chromosome sets (polyploids) are more stress- resistant and superior colonizers and may thus outnumber their low ploidy conspecifics in more extreme habitats. Brachypodium distachyon (sensu lato), for example, a common annual grass in Israel and the entire Mediterranean basin, comprises three cytotypes of differing chromosome numbers that were recently proposed as distinct species. It was suggested that increased aridity increases the occurrence of its polyploid cytotype. Here, we tested at two spatial scales whether polyploid plants of B. distachyon s. l. are more frequently found in drier habitats in Israel. We collected a total of 430 specimens (i) along a largescale climatic gradient with 15 thoroughly selected sites (spanning 114- 954 mm annual rainfall), and (ii) from corresponding Northern (more mesic) and Southern (more arid) hill slopes to assess the micro- climatic difference between contrasting exposures. Cytotypes were then determined via flow cytometry. Polyploid plants comprised 90% of all specimens and their proportion ranged between 0% and 100% per site. However, this proportion was not correlated with aridity along the large- scale gradient, nor were polyploids more frequently found on Southern exposures. Our results show for both spatial scales that increasing aridity is not the principal driver for the distribution of polyploids in B. distachyon s. l. in Israel. Notably, though, diploid plants were restricted essentially to four intermediate sites, while polyploids dominated the most arid and the most mesic sites. This, to some degree, clustered pattern suggests that the distribution of cytotypes is not entirely random and calls for future studies to assess further potential drivers.