Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

Contrasting mechanisms of secondary metabolite accumulation during leaf development in two tropical tree species with different leaf expansion strategies

Young leaves of most species experience remarkably higher herbivore attack rates than mature leaves. Considerable theoretical effort has focused on predicting optimal defense and tradeoffs in defense allocation during leaf expansion. Among others, allocation to secondary chemistry may be dependent on growth constraints. We studied flavanoid production during leaf development in two species of Inga (Fabaceae: Mimosoideae) with different expansion strategies: Inga goldmanii, a species with slowly expanding young leaves, and Inga umbellifera, a species with fast-expanding young leaves. In these two species, the most abundant and toxic class of defensive compounds is flavanoids (which include tannins). We measured their concentration by leaf dry weight, their total content per leaf, their HPLC chemical profile and their toxicity to a generalist herbivore at different expansion levels. Although in both species the flavanoid concentration decreased with increasing leaf expansion, that decrease was twice as pronounced for I. umbellifera as it was for I. goldmanii. I. umbellifera leaves produced flavanoids only during the first half of their development while I. goldmanii leaves continued production throughout. The changes in flavanoid HPLC profiles and toxicity were also more dramatic for I. umbellifera, which had different flavanoids in young than in mature leaves. Relative to I. umbellifera, I. goldmanii showed smaller changes in both flavanoid composition and toxicity in the transition from young to mature leaves. These results indicate that, even though young leaves suffer higher rates of attack and are predicted to have better chemical defenses than mature leaves, growth constraints may modulate defense allocation and thus, evolution of defense strategies.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Evaluation of leaf features in forest trees: Methods,techniques, obtainable information and limits

Tree leaves are interfaces between the whole organism and the environment. Leaves display a series of attributes that are linked to specific functions (functional leaf traits—FLT) and/or show responses to biotic and abiotic stress factors (stress response traits, SRT), which can be subdivided into: (a) morphological traits; (b) chemical traits; (c) physiological traits; (d) symptoms. The analysis of FLT is a useful tool for tree species and provenance phenotyping, due to the adaptation of trees to environmental stress. Additionally, FLT can be used as response factor in long term and large spatial scales surveys of forest conditions. Despite these potential benefits of leaf traits in the assessment of ecosystem health and functioning, leaf sampling in forests is time-consuming and costly, especially in forests with a complex vertical and horizontal structure and in remote forest areas. Once a foliar sample has been collected, many different analyses can be carried out; however, analyses should be technically simple and able to be performed within one day following the leaf collection (i.e., on fresh samples), or after air-drying the leaves themselves (analysis of dried specimens). This paper reports the results of leaf sampling and foliar analyses carried out in previous research projects and revises the current state-of-the-art. The leaf traits that are easily obtainable from leaf sampling are listed, together with the operational procedures necessary for their measurement, described in a standardized protocol. Their ecological and functional relevance is discussed in relation to their potential information (as indicators of climatic stress, drought, air and soil pollution, tree light-use and competition, plant nutritional status, health and general plant stress conditions). Finally, this review provides suggestions for the elaboration and reporting of data, and proposes some guidelines to improve the effectiveness of foliar analysis in the assessment of forest ecosystem health, properties and functioning.     

Physical,but not chemical,antiherbivore defense expression is related to the clustered spatial distribution of tropical trees in an Amazonian forest

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构树叶性状分化的生态功能初步研究

叶性状分化在自然界中较为普遍,不同的叶性状特征与植物对资源获得及利用效率密切关联,反映了植物适应特定环境所形成的生存对策。叶性状分化的生态功能一直以来备受生态学家和进化生物学家的广泛关注。自然界构树(Broussonetia papyifera)在个体发育过程中出现全缘叶和裂缺叶的分化,但其生态功能尚不清楚,推测两者的叶型分化是构树对虫害规避的结果。为了探讨构树叶性状分化对应的可能生态功能,该研究采用野外监测和室内分析的方法,对构树全缘叶和裂缺叶的虫害发生率、叶面积、与抗虫有关的酚类物质(总酚、缩合单宁、黄酮)含量进行了比较。结果表明:(1)相对于裂缺叶,全缘叶虫害发生率显著增加,全缘叶虫害发生率是裂缺叶的两倍。(2)自然条件下,全缘叶叶面积显著高于裂缺叶,增加了约44个百分点。(3)自然条件下,裂缺叶中总酚、缩合单宁、黄酮含量均显著高于全缘叶,分别提高了6.0%、4.2%和16.2%。(4)除黄酮外,虫害处理下裂缺叶中总酚、缩合单宁含量显著高于全缘叶,均提高了约5.0%。(5)人为移除部分叶片,裂缺叶中总酚、缩合单宁、黄酮含量均显著高于全缘叶,分别提高8.0%、1.6%和25.4%。这说明构树全缘叶和裂缺叶中酚类物质含量对外来损伤响应不一致,裂缺叶虫害发生率较全缘叶低可能由于两种类型叶片中酚类物质含量存在差异所引起。     

Unveiling the spatial distribution of aflatoxin B1 and plant defense metabolites in maize using AP-SMALDI mass spectrometry imaging

In order to cope with the presence of unfavorable compounds, plants can biotransform xenobiotics, translocate both parent compounds and metabolites, and perform compartmentation and segregation at the cellular or tissue level. Such a scenario also applies to mycotoxins, fungal secondary metabolites with a pre-eminent role in plant infection. In this work, we aimed to describe the effect of the interplay between Zea mays (maize) and aflatoxin B1 (AFB1) at the tissue and organ level. To address this challenge, we used atmospheric pressure scanning microprobe matrix-assisted laser desorption/ionization mass spectrometry imaging (AP-SMALDI MSI) to investigate the biotransformation, localization and subsequent effects of AFB1 on primary and secondary metabolism of healthy maize plants, both in situ and from a metabolomics standpoint. High spatial resolution (5 µm) provided fine localization of AFB1, which was located within the root intercellular spaces, and co-localized with its phase-I metabolite aflatoxin M2. We provided a parallel visualization of maize metabolic changes, induced in different organs and tissues by an accumulation of AFB1. According to our untargeted metabolomics investigation, anthocyanin biosynthesis and chlorophyll metabolism in roots are most affected. The biosynthesis of these metabolites appears to be inhibited by AFB1 accumulation. On the other hand, metabolites found in above-ground organs suggest that the presence of AFB1 may also activate the biochemical response in the absence of an actual fungal infection; indeed, several plant secondary metabolites known for their antimicrobial or antioxidant activities were localized in the outer tissues, such as phenylpropanoids, benzoxazinoids, phytohormones and lipids.     

Ants inhabiting myrmecophytic ferns regulate the distribution of lianas on emergent trees in a Bornean tropical rainforest

Little is known about the spatial distribution of lianas on emergent trees in tropical rainforests and the factors affecting this distribution. The present study investigated the effects of an arboreal ant species, Crematogaster difformis, which forms myrmecophytic symbioses with two epiphytic ferns, Lecanopteris sp. and Platycerium sp., on the spatial distribution of lianas associated with emergent trees. Living lianas were placed onto trunk surfaces inside and outside the territories of the ants in the canopy, to examine their ability to remove them. The number of leaves pruned by the ants was significantly higher on lianas inside than outside their territories. The spatial overlap of the distributions of lianas and the two ferns on emergent trees were then examined. The frequency of liana colonization of tree crowns was found to be significantly lower on trees with than without ferns. Under the natural conditions, C. difformis workers were observed biting and pruning the lianas. These results suggest that C. difformis regulates the distribution of lianas on emergent trees.     

Use of an exotic host plant shifts immunity,chemical defense,and viral burden in wild populations of a specialist insect herbivore

Defense against natural enemies constitutes an important driver of herbivore host range evolution in the wild. Populations of the Baltimore checkerspot butterfly, Euphydryas phaeton (Nymphalidae), have recently incorporated an exotic plant, Plantago lanceolata (Plantaginaceae), into their dietary range. To understand the tritrophic consequences of utilizing this exotic host plant, we examined immune performance, chemical defense, and interactions with a natural entomopathogen (Junonia coenia densovirus, Parvoviridae) across wild populations of this specialist herbivore. We measured three immune parameters, sequestration of defensive iridoid glycosides (IGs), and viral infection load in field‐collected caterpillars using either P. lanceolata or a native plant, Chelone glabra (Plantaginaceae). We found that larvae using the exotic plant exhibited reduced immunocompetence, compositional differences in IG sequestration, and higher in situ viral burdens compared to those using the native plant. On both host plants, high IG sequestration was associated with reduced hemocyte concentration in the larval hemolymph, providing the first evidence of incompatibility between sequestered chemical defenses and the immune response (i.e., the “vulnerable host” hypothesis) from a field‐based study. However, despite this negative relationship between IG sequestration and cellular immunity, caterpillars with greater sequestration harbored lower viral loads. While survival of virus‐infected individuals decreased with increasing viral burden, it ultimately did not differ between the exotic and native plants. These results provide evidence that: (1) phytochemical sequestration may contribute to defense against pathogens even when immunity is compromised and (2) herbivore persistence on exotic plant species may be facilitated by sequestration and its role in defense against natural enemies.     

Plant development controls leaf area expansion in alfalfa plants competing for light

Background and Aims

The growth of crops in a mixture is more variable and difficult to predict than that in pure stands. Light partitioning and crop leaf area expansion play prominent roles in explaining this variability. However, in many crops commonly grown in mixtures, including the forage species alfalfa, the sensitivity and relative importance of the physiological responses involved in the light modulation of leaf area expansion are still to be established. This study was designed to assess the relative sensitivity of primary shoot development, branching and individual leaf expansion in alfalfa in response to light availability.

Methods

Two experiments were carried out. The first studied isolated plants to assess the potential development of different shoot types and growth periods. The second consisted of manipulating the intensity of competition for light using a range of canopies in pure and mixed stands at two densities so as to evaluate the relative effects on shoot development, leaf growth, and plant and shoot demography.

Key Results

Shoot development in the absence of light competition was deterministic (constant phyllochrons of 32·5 °Cd and 48·2 °Cd for primary axes and branches, branching probability of 1, constant delay of 1·75 phyllochron before axillary bud burst) and identical irrespective of shoot type and growth/regrowth periods. During light competition experiments, changes in plant development explained most of the plant leaf area variations, with average leaf size contributing to a lesser extent. Branch development and the number of shoots per plant were the leaf area components most affected by light availability. Primary axis development and plant demography were only affected in situations of severe light competition.

Conclusions

Plant leaf area components differed with regard to their sensitivity to light competition. The potential shoot development model presented in this study could serve as a framework to integrate light responses in alfalfa crop models.     

Contributions of leaf photosynthetic capacity, leaf angle and self-shading to the maximization of net photosynthesis in Acer saccharum: a modelling assessment

Background and Aims

Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ɛ).

Methods

A functional–structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI_ind) together with a genetic algorithm to find distributions of leaf angle (L_A) and leaf photosynthetic capacity (A_max) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A_max either unconstrained or constrained to an upper value consistent with reported values for A_max in A. saccharum.

Key Results

It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ɛ were simultaneously maximized. Maximization of ɛ required simultaneous adjustments in L_A and A_max along gradients of PPFD in the plants. When A_max was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ɛ because PPFD incident on leaves was higher than the PPFD at which ɛ_max was attainable. Average leaf ɛ in constrained plants nonetheless improved with increasing LAI_ind because of an increase in self-shading.

Conclusions

It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ɛ at the scale of leaves, which requires a highly integrated response between L_A, A_max and LAI_ind. The results also suggest that to maximize ɛ plants have evolved mechanisms that co-ordinate the L_A and A_max of individual leaves with PPFD availability.     

Effects of insect herbivory on induced chemical defences and compensation during early plant development in Penstemon virgatus

Background and Aims

The lack of studies assessing the simultaneous expression of tolerance and resistance traits during seedling development and overall seedling defences as compared with adult plants, in general, constitutes a significant research need that can greatly improve our understanding of overall investment in defences during plant ontogeny.

Methods

Using two seedling and two juvenile stages of the perennial herb Penstemon virgatus (Plantaginaceae) evaluations were made of (a) patterns of investment in constitutive chemical defences [i.e. iridoid glycosides (IGs)], and (b) simultaneous variation in the short-term ability of seedling and juvenile stages to induce resistance traits, measured as induced chemical defences, or tolerance traits, measured as compensatory re-growth following moderate levels of damage by a specialist insect herbivore.

Key Results

Plants were highly defended during most of their transition from seedling to early juvenile stages, reaching a constant approx. 20 % dry weight total IGs. Furthermore, following 30 % above-ground tissue damage, seedlings and juvenile stages were equally able to induce resistance, by raising their IG concentration by approx. 8 %, whereas compensatory re-growth was only achieved at young juvenile but not seedling stages.

Conclusions

Two major trends emerged from this study: (1) in contrast to expected and previously observed trends, in this perennial plant species, seedlings seem to be one of the most well-defended stages as compared with adult ones; (2) high levels of constitutive defences did not limit the ability of young developmental stages to induce resistance following damage, although this response may come with a cost (i.e. decreased compensation) in young seedling stages. Hence, as has been previously demonstrated in few other systems, these results points towards an indirect evidence for a trade-off between tolerance and resistance traits at some, but not all, developmental stages; making them often difficult to detect.     

wing blister, a new Drosophila laminin alpha chain required for cell adhesion and migration during embryonic and imaginal development

We report the molecular and functional characterization of a new alpha chain of laminin in Drosophila. The new laminin chain appears to be the Drosophila counterpart of both vertebrate alpha2 (also called merosin) and alpha1 chains, with a slightly higher degree of homology to alpha2, suggesting that this chain is an ancestral version of both alpha1 and alpha2 chains. During embryogenesis, the protein is associated with basement membranes of the digestive system and muscle attachment sites, and during larval stage it is found in a specific pattern in wing and eye discs. The gene is assigned to a locus called wing blister (wb), which is essential for embryonic viability. Embryonic phenotypes include twisted germbands and fewer pericardial cells, resulting in gaps in the presumptive heart and tracheal trunks, and myotubes detached from their target muscle attachment sites. Most phenotypes are in common with those observed in Drosophila laminin alpha3, 5 mutant embryos and many are in common with those observed in integrin mutations. Adult phenotypes show blisters in the wings in viable allelic combinations, similar to phenotypes observed in integrin genes. Mutation analysis in the eye demonstrates a function in rhabdomere organization. In summary, this new laminin alpha chain is essential for embryonic viability and is involved in processes requiring cell migration and cell adhesion.     

Linking ethylene to nitrogen-dependent leaf longevity of grass species in a temperate steppe

Background and Aims

Leaf longevity is an important plant functional trait that often varies with soil nitrogen supply. Ethylene is a classical plant hormone involved in the control of senescence and abscission, but its role in nitrogen-dependent leaf longevity is largely unknown.

Methods

Pot and field experiments were performed to examine the effects of nitrogen addition on leaf longevity and ethylene production in two dominant plant species, Agropyron cristatum and Stipa krylovii, in a temperate steppe in northern China.

Key Results

Nitrogen addition increased leaf ethylene production and nitrogen concentration but shortened leaf longevity; the addition of cobalt chloride, an ethylene biosynthesis inhibitor, reduced leaf nitrogen concentration and increased leaf longevity. Path analysis indicated that nitrogen addition reduced leaf longevity mainly through altering leaf ethylene production.

Conclusions

These findings provide the first experimental evidence in support of the involvement of ethylene in nitrogen-induced decrease in leaf longevity.     

性别和发育阶段对绒毛白蜡光合特征及叶功能性状的影响

雌雄异株植物资源分配模式上往往表现出显著的性别二态性,但在叶片光合及功能性状上是否有差异目前仍未有定论,且与发育阶段的关系尚不明确。阐明上述问题,能够进一步了解雌雄异株植物的生理生态特征,并为理解性别对性二态植物生长发育的影响机制提供理论依据。以雌雄异株绒毛白蜡(Fraxinus velutina Torr.)为研究对象,针对不同发育阶段不同性别植株进行光合特征及叶功能性状测定,采用双因素方差分析了不同发育阶段下雌雄植株光合能力及叶功能性状的性别间差异,采用Pearson检验了雌雄植株各叶功能性状之间的相关性,并采用标准化主轴分析(Standardized major axis regression, SMA)分析不同性别植株净光合速率与叶功能性状的相关性。结果表明性别和发育阶段显著影响植物个体的光合能力和叶功能性状。总体而言,雄树在坐果期和果实成熟期均表现出更强的净光合速率(Pn)、更高的比叶面积(SLA)、叶绿素含量(Chl)和叶氮含量(LNC);而雌树在果实膨大期表现出更强Pn、SLA和Chl。雌雄性别内Pn与SLA、Chl和LNC间均呈显著正相关(P<0.05),雄树的S...     

Correlation of Versican Expression,Accumulation, and Degradation during Embryonic Development by Quantitative Immunohistochemistry

Versican, a chondroitin sulfate proteoglycan, is important in embryonic development, and disruption of the versican gene is embryonically lethal in the mouse. Although several studies show that versican is increased in various organs during development, a focused quantitative study on versican expression and distribution during lung and central nervous system development in the mouse has not previously been performed. We tracked changes in versican (Vcan) gene expression and in the accumulation and degradation of versican. Vcan expression and quantitative immunohistochemistry performed from embryonic day (E) 11.5 to E15.5 showed peak Vcan expression at E13.5 in the lungs and brain. Quantitative mRNA analysis and versican immunohistochemistry showed differences in the expression of the versican isoforms in the embryonic lung and head. The expression of Vcan mRNA and accumulation of versican in tissues was complementary. Immunohistochemistry demonstrated co-localization of versican accumulation and degradation, suggesting distinct roles of versican deposition and degradation in embryogenesis. Very little versican mRNA or protein was found in the lungs of 12- to 16-week-old mice but versican accumulation was significantly increased in mice with Pseudomonas aeruginosa lung infection. These data suggest that versican plays an important role in fundamental, overlapping cellular processes in lung development and infection.     

Effects of light quality on leaf morphogenesis of a heterophyllous amphibious plant, Rotala hippuris

Background and Aims

For heterophyllous amphibious plants that experience fluctuating water levels, it is critical to control leaf development precisely in response to environmental cues that can serve as a quantitative index of water depth. Light quality can serve as such a cue because the ratio of red light relative to far-red light (R/FR) increases and blue-light intensity decreases with increasing water depth. Growth experiments were conducted to examine how R/FR and blue-light intensity alter leaf morphology of a heterophyllous amphibious plant, Rotala hippuris.

Methods

Using combinations of far red (730 nm), red (660 nm) and blue (470 nm) light-emitting diodes (LEDs), growth experiments were used to quantitatively evaluate the effects of the R/FR ratio and blue-light intensity on leaf morphology.

Key Results

Under the natural light regime in an outside growth garden, R. hippuris produced distinct leaves under submerged and aerial conditions. R/FR and blue-light intensity were found to markedly affect heterophyllous leaf formation. Higher and lower R/FR caused leaf characters more typical of submerged and aerial leaves, respectively, in both aerial and submerged conditions, in accordance with natural distribution of leaf types and light under water. High blue light caused a shift of trait values toward those of typical aerial leaves, and the response was most prominent under conditions of R/FR that were expected near the water surface.

Conclusions

R/FR and blue-light intensity provides quantitative cues for R. hippuris to detect water depth and determine the developmental fates of leaves, especially near the water surface. The utilization of these quantitative cues is expected to be important in habitats where plants experience water-level fluctuation.