Are correlations among foliar traits in ferns consistent with those in the seed plants?

Broad-based studies of gymnosperms and angiosperms reveal consistent and functionally significant correlations among foliar traits such as leaf mass per area (LMA), maximum photosynthetic rate (A(area)), foliar nitrogen (N(area)), foliar chlorophyll (Chl) and leaf longevity. To assess the generality of these relationships, we studied 20 fern species growing in the understorey of a temperate deciduous forest. We found that foliar N(area) increases with LMA, and that foliar N(area) and A(area) are positively correlated with one another, as are foliar N(area) and Chl. The ferns in general have very low LMA compared with most seed plants; A(area), N(area) and Chl are below median values for seed plants but are not extreme. Species with overwintering fronds have significantly higher LMA than species with fronds that senesce at the end of the growing season, as well as a significantly higher C : N ratio in frond tissue and relatively high foliar N on an areal basis. Correlations among foliar traits associated with gas exchange in these forest understorey ferns are in accordance with patterns reported for seed plants, suggesting a high degree of functional constraint on the interrelationships among key elements in foliar design.     

Changes in hydraulic traits of nine vascular plants from different evolutionary lineages北大核心CSCD

Aims As vascular plants evolve from ferns to gymnosperms and angiosperms, their physiological structures and functions are assumed more adaptable to arid environment. Whether the three plant groups from early to late evolved lineages have improved their water transport and use efficiency has been studied on the basis of the morphological structure of leaf veins and stomata.Moreover, the water transportation rate was directly measured in the angiosperms. Therefore, we measured structural and functional traits related to water relations in all three plant groups simultaneously, to test the hypothesis on the evolutionary process of plant hydraulics. Methods We selected three species in each group grown in South China Botanical Garden, Guangzhou, China, including ferns (Dicranopteris pedata, Cyclosorus parasiticus and Blechnum orientale), gymnosperms (Podocarpus macrophyllus, Podocarpus nagi and Taxodium distichum) and angiosperms (Manglietia fordiana var. hainanensis, Sindora tonkinensis and Bauhinia purpurea). Important findings Sapwood and leaf specific hydraulic conductivities (KS and KL, respectively), and leaf conductance (Kleaf) significantly increased from ferns, gymnosperms to angiosperms. However, no significant trends were found in transpiration rate (E) and intrinsic water use efficiency. Meanwhile, neither the size and density of stomata nor wood density showed significant difference among three plant groups. The hydraulic functional traits (KS, KL and Kleaf) had significantly positive correlations with each other, but had no relationships with the two measured structural traits. Phylogenetic independent contrasts analyses showed that the coordination between KS and Kleaf, and between KS and E were independent of the phylogeny. Based on the nine vascular species, this study demonstrated that water transport related traits are improved as vascular plants evolved, and the co-evolution between water transport and transpiration traits were identified. For further study, it is necessary to consolidate our data with investigations of more detailed water-transport structures in more species from different evolutionary lineages. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Contrasting Predictors of Fern versus Angiosperm Decomposition in a Common Garden

We evaluated differences in the rates and correlates of decomposition among 32 fern and angiosperm litter types collected in Hawai'i. Leptosporangiate ferns were separated into groups based on phylogeny: 'polypod' ferns, a monophyletic clade of ferns that diversified in the Cretaceous, and all other ('non-polypod') ferns that diversified earlier. We measured initial litter chemistry (nutrients and carbon chemistry), and mass loss and nitrogen (N), phosphorus (P), and calcium (Ca) of litter tissue during a 1-yr incubation in a common garden. Nutrient concentrations and carbon (C) chemistry differed significantly among litter types, and litter turnover ( k -values) ranged from 0.29 to 8.31. Decomposition rates were more closely correlated with nutrient concentration than is typically observed. Lignin:N was the best predictor of decomposition across all litter types combined; however, among plant groups different predictors of decomposition were important. Nitrogen and P concentrations best predicted fern decomposition, whereas C chemistry, particularly lignin concentration, was more important for angiosperm (monocot and dicot) decomposition. Among native plants, non-polypod ferns decomposed significantly more slowly than both polypod ferns and angiosperms. Contrary to our hypothesis, fern litter did not decompose more slowly than angiosperm litter overall. Nutrient dynamics in litter were affected by initial litter concentration more than phylogeny; low-nutrient litter immobilized more nutrients than high-nutrient litter. Systematic differences in rates of decomposition, and the importance of nutrients in predicting fern decomposition, imply that changes in species composition within ferns and between ferns and angiosperms could influence the functioning of ecosystems where ferns are important forest components.     

KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems

Members of the class 1 knotted-like homeobox (KNOX) gene family are important regulators of shoot apical meristem development in angiosperms. To determine whether they function similarly in seedless plants, three KNOX genes (two class 1 genes and one class 2 gene) from the fern Ceratopteris richardii were characterized. Expression of both class 1 genes was detected in the shoot apical cell, leaf primordia, marginal part of the leaves, and vascular bundles by in situ hybridization, a pattern that closely resembles that of class 1 KNOX genes in angiosperms with compound leaves. The fern class 2 gene was expressed in all sporophyte tissues examined, which is characteristic of class 2 gene expression in angiosperms. All three CRKNOX genes were not detected in gametophyte tissues by RNA gel blot analysis. Arabidopsis plants overexpressing the fern class 1 genes resembled plants that overexpress seed plant class 1 KNOX genes in leaf morphology. Ectopic expression of the class 2 gene in Arabidopsis did not result in any unusual phenotypes. Taken together with phylogenetic analysis, our results suggest that (a) the class 1 and 2 KNOX genes diverged prior to the divergence of fern and seed plant lineages, (b) the class 1 KNOX genes function similarly in seed plant and fern sporophyte meristem development despite their differences in structure, (c) KNOX gene expression is not required for the development of the fern gametophyte, and (d) the sporophyte and gametophyte meristems of ferns are not regulated by the same developmental mechanisms at the molecular level.     

九种维管植物水力性状的演化趋势

维管植物从蕨类植物、裸子植物到被子植物的演化过程中, 生理结构和功能都被认为更趋向于适应干旱的环境。但是关于3类植物是否提升了水分传导和水分利用效率的研究目前多基于叶脉和气孔的形态结构推断, 而对水分传导速率的直接测量则集中于被子植物内部。因此, 同时测定3类植物叶片和枝条尺度的水力性状, 可以研究维管植物水力性状的演化趋势和协同演化关系。该研究测定了中国科学院华南植物园内的蕨类植物(芒萁(Dicranopteris pedata)、华南毛蕨(Cyclosorus parasiticus)、乌毛蕨(Blechnum orientale)), 裸子植物(罗汉松(Podocarpus macrophyllus)、竹柏(Podocarpus nagi)、落羽杉(Taxodium distichum))和被子植物(海南木莲(Manglietia fordiana var. hainanensis)、东京油楠(Sindora tonkinensis)、羊蹄甲(Bauhinia purpurea))共9种植物的水力结构与功能的相关性状。发现9种植物从原始到更演化物种的枝条比导水率(K_S)、叶片比导水率(K_L)和叶片水力导度(K_leaf)依次升高; 叶片蒸腾速率(E)和潜在水分利用效率(WUE_i)升高趋势不显著; 气孔面积指数(SPI)和木材密度(WD)在3个类群间无明显变化趋势。水力功能性状在枝条(K_S)和叶片(K_L、K_leaf和E)尺度显著相关, 但和两个结构性状并无显著相关性。对显著相关的性状进行系统发育独立差分析发现相关性依然存在, 表明枝条导水和叶片导水、叶片失水的协同性独立于系统发育而存在。该文以9种维管植物为例, 发现演化过程中水分传导相关的功能性状有显著提升, 且导水和失水的性状存在着协同演化关系。上述维管植物水力性状演化趋势的结论需要对更多物种、更精细的水力结构性状的进行测定验证。     

Changes in hydraulic traits of nine vascular plants from different evolutionary lineages

维管植物从蕨类植物、裸子植物到被子植物的演化过程中, 生理结构和功能都被认为更趋向于适应干旱的环境。但是关于3类植物是否提升了水分传导和水分利用效率的研究目前多基于叶脉和气孔的形态结构推断, 而对水分传导速率的直接测量则集中于被子植物内部。因此, 同时测定3类植物叶片和枝条尺度的水力性状, 可以研究维管植物水力性状的演化趋势和协同演化关系。该研究测定了中国科学院华南植物园内的蕨类植物(芒萁(Dicranopteris pedata)、华南毛蕨(Cyclosorus parasiticus)、乌毛蕨(Blechnum orientale)), 裸子植物(罗汉松(Podocarpus macrophyllus)、竹柏(Podocarpus nagi)、落羽杉(Taxodium distichum))和被子植物(海南木莲(Manglietia fordiana var. hainanensis)、东京油楠(Sindora tonkinensis)、羊蹄甲(Bauhinia purpurea))共9种植物的水力结构与功能的相关性状。发现9种植物从原始到更演化物种的枝条比导水率(K_S)、叶片比导水率(K_L)和叶片水力导度(K_leaf)依次升高; 叶片蒸腾速率(E)和潜在水分利用效率(WUE_i)升高趋势不显著; 气孔面积指数(SPI)和木材密度(WD)在3个类群间无明显变化趋势。水力功能性状在枝条(K_S)和叶片(K_L、K_leaf和E)尺度显著相关, 但和两个结构性状并无显著相关性。对显著相关的性状进行系统发育独立差分析发现相关性依然存在, 表明枝条导水和叶片导水、叶片失水的协同性独立于系统发育而存在。该文以9种维管植物为例, 发现演化过程中水分传导相关的功能性状有显著提升, 且导水和失水的性状存在着协同演化关系。上述维管植物水力性状演化趋势的结论需要对更多物种、更精细的水力结构性状的进行测定验证。     

A mechanism for the leaching of calcium from foliage

Young bean plants (Phaseolus vulgaris) containing root-absorbed ⁴⁵Ca and ⁸⁶Rb were leached to determine the pathway and mechanism of cation loss by leaching. Calcium is leached from the exchangeable calcium fraction within the plant by a process of ion exchange and diffusion involving exchange sites both within the leaf and on the leaf surface. Leaching of cations is primarily a passive process, although some metabolites may be deposited upon leaf surfaces by active processes. The exchange and diffusion explanation is compatible with current theories of ion uptake and translocation and explains the results of numerous experiments on leaching reported in the literature.     

Fern and lycophyte guard cells do not respond to endogenous abscisic acid

Stomatal guard cells regulate plant photosynthesis and transpiration. Central to the control of seed plant stomatal movement is the phytohormone abscisic acid (ABA); however, differences in the sensitivity of guard cells to this ubiquitous chemical have been reported across land plant lineages. Using a phylogenetic approach to investigate guard cell control, we examined the diversity of stomatal responses to endogenous ABA and leaf water potential during water stress. We show that although all species respond similarly to leaf water deficit in terms of enhanced levels of ABA and closed stomata, the function of fern and lycophyte stomata diverged strongly from seed plant species upon rehydration. When instantaneously rehydrated from a water-stressed state, fern and lycophyte stomata rapidly reopened to predrought levels despite the high levels of endogenous ABA in the leaf. In seed plants under the same conditions, high levels of ABA in the leaf prevented rapid reopening of stomata. We conclude that endogenous ABA synthesized by ferns and lycophytes plays little role in the regulation of transpiration, with stomata passively responsive to leaf water potential. These results support a gradualistic model of stomatal control evolution, offering opportunities for molecular and guard cell biochemical studies to gain further insights into stomatal control.     

An exceptional role for flowering plant physiology in the expansion of tropical rainforests and biodiversity

Movement of water from soil to atmosphere by plant transpiration can feed precipitation, but is limited by the hydraulic capacities of plants, which have not been uniform through time. The flowering plants that dominate modern vegetation possess transpiration capacities that are dramatically higher than any other plants, living or extinct. Transpiration operates at the level of the leaf, however, and how the impact of this physiological revolution scales up to the landscape and larger environment remains unclear. Here, climate modelling demonstrates that angiosperms help ensure aseasonally high levels of precipitation in the modern tropics. Most strikingly, replacement of angiosperm with non-angiosperm vegetation would result in a hotter, drier and more seasonal Amazon basin, decreasing the overall area of ever-wet rainforest by 80 per cent. Thus, flowering plant ecological dominance has strongly altered climate and the global hydrological cycle. Because tropical biodiversity is closely tied to precipitation and rainforest area, angiosperm climate modification may have promoted diversification of the angiosperms themselves, as well as radiations of diverse vertebrate and invertebrate animal lineages and of epiphytic plants. Their exceptional potential for environmental modification may have contributed to divergent responses to similar climates and global perturbations, like mass extinctions, before and after angiosperm evolution.     

The invention of WUS-like stem cell-promoting functions in plants predates leptosporangiate ferns

The growth of land plants depends on stem cell-containing meristems which show major differences in their architecture from basal to higher plant species. In Arabidopsis, the stem cell niches in the shoot and root meristems are promoted by WUSCHEL (WUS) and WOX5, respectively. Both genes are members of a non-ancestral clade of the WUS-related homeobox (WOX) gene family, which is absent in extant bryophytes and lycophytes. Our analyses of five fern species suggest that a single WUS orthologue was present in the last common ancestor (LCA) of leptosporangiate ferns and seed plants. In the extant fern Ceratopteris richardii, the WUS pro-orthologue marks the pluripotent cell fate of immediate descendants of the root apical initial, so-called merophytes, which undergo a series of stereotypic cell divisions and give rise to all cell types of the root except the root cap. The invention of a WUS-like function within the WOX gene family in an ancestor of leptosporangiate ferns and seed plants and its amplification and sub-functionalisation to different stem cell niches might relate to the success of seed plants, especially angiosperms.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

蕨类植物配子体育及其性器官分化的研究进展

作者概述了蕨类植物生活周期中不同发育阶段的特点,并着重介绍了不,植物生长调节物质,水和温度等因素对蕨类植物配子体发育的影响以及精子囊素,光,钙离子和其它植物生长调节物质在性器官形成和分化过程中的作用,同时作者对蕨类植物性器官分化的不稳定性进行了探讨,并结合目前蕨类植物性器官分化的遗传学和分子生物学研究的最新进展,说明蕨类植物作为研究系统在植物生物学研究领域中的优势。     

蕨类植物配子体发育及其性器官分化的研究进展

作者概述了蕨类植物生活周期中不同发育阶段的特点,并着重介绍了光、植物生长调节物质、水和温度等因素对蕨类植物配子体发育的影响以及成精子囊素、光、钙离子和其它植物生长调节物质在性器官形成和分化过程中的作用。同时作者对蕨类植物性器官分化的不稳定性进行了探讨、并结合目前蕨类植物性器官分化的遗传学和分子生物学研究的最新进展,说明蕨类植物作为研究系统在植物生物学研究领域中的优势。     

海南和台湾蕨类植物多样性及其大陆性特征

海南和台湾是我国南部和东部两个最重要的大陆性岛屿,具有极为丰富的蕨类植物多样性。特有现象揭示着植物区系和植物多样性的历史,间断分布揭示着与邻近或相关植物区系的联系;特别是在岛屿地区,这种现象和意义尤其明显。海南有蕨类植物区系55科、135属、466种,其中特有种有32种,台湾蕨类植物区系57科、142属、599种,其中特有种达66种。在海南与台湾两地,有共有属104属,共有种仅有176种。台湾海峡出现始于晚白垩世,持续至第四纪;琼州海峡出现于早第四纪,因此,两地的特有现象远少于种子植物(台湾有801种,海南有501种),原因与蕨类植物具有更广的散布性相关,而且在被子植物中起重要作用的物种生物学障碍(机制)在蕨类是缺乏的;形态学的、生殖生物学特征导致蕨类植物具有较缓慢的物种演化历史和较低的灭绝率(Smith,1972)。海南和台湾蕨类植物区系的比较还表明,地理位置和海拔高度对植物区系的物种分化和物种多样性产生了极大的影响。     

Phylogenetic variation in the silicon composition of plants

BACKGROUND AND AIMS: Silicon (Si) in plants provides structural support and improves tolerance to diseases, drought and metal toxicity. Shoot Si concentrations are generally considered to be greater in monocotyledonous than in non-monocot plant species. The phylogenetic variation in the shoot Si concentration of plants reported in the primary literature has been quantified. METHODS: Studies were identified which reported Si concentrations in leaf or non-woody shoot tissues from at least two plant species growing in the same environment. Each study contained at least one species in common with another study. KEY RESULTS: Meta-analysis of the data revealed that, in general, ferns, gymnosperms and angiosperms accumulated less Si in their shoots than non-vascular plant species and horsetails. Within angiosperms and ferns, differences in shoot Si concentration between species grouped by their higher-level phylogenetic position were identified. Within the angiosperms, species from the commelinoid monocot orders Poales and Arecales accumulated substantially more Si in their shoots than species from other monocot clades. CONCLUSIONS: A high shoot Si concentration is not a general feature of monocot species. Information on the phylogenetic variation in shoot Si concentration may provide useful palaeoecological and archaeological information, and inform studies of the biogeochemical cycling of Si and those of the molecular genetics of Si uptake and transport in plants.     

Leaf energy balance modelling as a tool to infer habitat preference in the early angiosperms

Despite more than a century of research, some key aspects of habitat preference and ecology of the earliest angiosperms remain poorly constrained. Proposed growth ecology has varied from opportunistic weedy species growing in full sun to slow-growing species limited to the shaded understorey of gymnosperm forests. Evidence suggests that the earliest angiosperms possessed low transpiration rates: gas exchange rates for extant basal angiosperms are low, as are the reconstructed gas exchange rates for the oldest known angiosperm leaf fossils. Leaves with low transpirational capacity are vulnerable to overheating in full sun, favouring the hypothesis that early angiosperms were limited to the shaded understorey. Here, modelled leaf temperatures are used to examine the thermal tolerance of some of the earliest angiosperms. Our results indicate that small leaf size could have mitigated the low transpirational cooling capacity of many early angiosperms, enabling many species to survive in full sun. We propose that during the earliest phases of the angiosperm leaf record, angiosperms may not have been limited to the understorey, and that some species were able to compete with ferns and gymnosperms in both shaded and sunny habitats, especially in the absence of competition from more rapidly growing and transpiring advanced lineages of angiosperms.     

Recurrent genome duplication events likely contributed to both the ancient and recent rise of ferns

Ferns, the second largest group of vascular plants, originated ～400 mil ion years ago(Mya). They became dominant in the ancient Earth landscape before the angiosperms and are stil important in current ecosystems.Many ferns have exceptional y high chromosome numbers,possibly resulting from whole-genome duplications(WGDs).However, WGDs have not been investigated molecularly across fern diversity. Here we detected and dated fern WGDs using a phylogenomic approach and by calculating synonymous substitution rates(Ks). We also investigated a possible correlation between proposed WGDs and shifts in species diversification rates. We identified 19 WGDs: three ancient events along the fern phylogenetic backbone that are shared by 66%–97% of extant ferns, with additional lineage-specific WGDs for eight orders, providing strongevidence for recurring genome duplications across fern evolutionary history. We also observed similar Ks peak values for more than half of these WGDs, with multiple WGDs occurring close to the Cretaceous(～145–66 Mya). Despite the repeated WGD events, the biodiversity of ferns declined during the Cretaceous, implying that other factors probably contributed to the floristic turnover from ferns to angiosperms. This study provides molecular evidence for recurring WGDs in ferns and offers important clues to the genomic evolutionary history of ferns.     

Diffusional limitations explain the lower photosynthetic capacity of ferns as compared with angiosperms in a common garden study

Ferns are thought to have lower photosynthetic rates than angiosperms and they lack fine stomatal regulation. However, no study has directly compared photosynthesis in plants of both groups grown under optimal conditions in a common environment. We present a common garden comparison of seven angiosperms and seven ferns paired by habitat preference, with the aims of (1) confirming that ferns do have lower photosynthesis capacity than angiosperms and quantifying these differences; (2) determining the importance of diffusional versus biochemical limitations; and (3) analysing the potential implication of leaf anatomical traits in setting the photosynthesis capacity in both groups. On average, the photosynthetic rate of ferns was about half that of angiosperms, and they exhibited lower stomatal and mesophyll conductance to CO₂ (g_m), maximum velocity of carboxylation and electron transport rate. A quantitative limitation analysis revealed that stomatal and mesophyll conductances were co‐responsible for the lower photosynthesis of ferns as compared with angiosperms. However, g_m alone was the most constraining factor for photosynthesis in ferns. Consistently, leaf anatomy showed important differences between angiosperms and ferns, especially in cell wall thickness and the surface of chloroplasts exposed to intercellular air spaces.     

Cross-phytogroup assessment of foliar epiphytic mycobiomes

The foliar surface forms one of the largest aboveground habitats on Earth and maintains plant-fungus relationships that greatly affect ecosystem functioning. Despite many studies with particular plant species, the foliar epiphytic mycobiome has not been studied across a large number of plant species from different taxa. Using high-throughput sequencing, we assessed epiphytic mycobiomes on leaf surfaces of 592 plant species in a botanical garden. Plants of angiosperms, gymnosperms, and pteridophytes were involved. Plant taxonomy, leaf side, growing environment, and evolutionary relationships were considered. We found that pteridophytes showed the higher fungal species diversity, stronger mutualistic fungal interactions, and a greater percentage of putative pathogens than gymnosperms and angiosperms. Plant taxonomic group, leaf side, and growing environment were significantly associated with the foliar epiphytic mycobiome, but the similarity of the mycobiomes among plants was not directly related to the distance of the host evolutionary tree. Our results provide a general understanding of the foliar fungal mycobiomes from pteridophytes to angiosperms. These findings will facilitate our understanding of foliar fungal epiphytes and their roles in plant communities and ecosystems.     

Leaf hydraulic capacity in ferns, conifers and angiosperms: impacts on photosynthetic maxima

* The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. * Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. * We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance (Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. * The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.