Does disturbance regime change community assembly of angiosperm plant communities in the boreal forest?

Aims To examine if and how species and phylogenetic diversity change in relation to disturbance, we conducted a review of ecological literature by testing the consistency of the relationship between phylogenetic diversity and disturbance and compared taxonomic groups, type of disturbance and ecosystem/habitat context. We provide a case study of the phylogenetic diversity–disturbance relationship in angiosperm plant communities of a boreal forest region, compared with types of natural and anthropogenic disturbances and plant growth forms.Methods Using a large-scale sampling plot network along a complete (0–100%) anthropogenic disturbance gradient in the boreal biome, we compared the changes of angiosperm plant community structure and composition across plots. We estimated natural disturbance with historical records of major fires. We then calculated phylogenetic diversity indexes and determined species richness in order to compare linear and polynomial trends along disturbance gradients. We also compared the changes of community structure for different types of anthropogenic disturbances and examined how the relationships between species and phylogenetic diversity and disturbance regimes vary among three different life forms (i.e. forbs, graminoids and woody plants).Important findings Phylogenetic diversity was inconsistently related to disturbance in previous studies, regardless of taxon, disturbance type or ecosystem context. In the understudied boreal ecosystem, angiosperm plant communities varied greatly in species richness and phylogenetic diversity along anthropogenic disturbance gradients and among different disturbance types. In general, a quadratic curve described the relationship between species richness and anthropogenic disturbance, with the highest richness at intermediate anthropogenic disturbance levels. However, phylogenetic diversity was not related to disturbance in any consistent manner and species richness was not correlated with phylogenetic diversity. Phylogenetic relatedness was also inconsistent across plant growth forms and different anthropogenic disturbance types. Unlike the inconsistent patterns observed for anthropogenic disturbance, community assembly among localities varying in time since natural disturbance exhibited a distinct signature of phylogenetic relatedness, although those trends varied among plant growth forms.     

Does forest fragmentation cause an increase in forest temperature?

Forest fragmentation is considered by many to be a primary cause of the current biodiversity crisis. The underlying mechanisms are poorly known, but a potentially important one is associated with altered thermal conditions within the remaining forest patches, especially at forest edges. Yet, large uncertainty remains about the effect of fragmentation on forest temperature, as it is unclear whether temperature decreases from forest edge to forest interior, and whether this local gradient scales up to an effect of fragmentation (landscape attribute) on temperature. We calculated the effect size (correlation coefficient) of distance from forest edge on air temperature, and tested for differences among forest types surrounded by different matrices using meta-analysis techniques. We found a negative edge-interior temperature gradient, but correlation coefficients were highly variable, and significant only for temperate and tropical forests surrounded by a highly contrasting open matrix. Nevertheless, it is unclear if these local-scale changes in temperature can be scaled up to an effect of fragmentation on temperature. Although it may be valid when considering “fragmentation” as forest loss only, the landscape-scale inference is not so clear when we consider the second aspect of fragmentation, where a given amount of forest is divided into a large number of small patches (fragmentation per se). Therefore, care is needed when assuming that fragmentation changes forest temperature, as thermal changes at forest edges depend on forest type and matrix composition, and it is still uncertain if this local gradient can be scaled up to the landscape.     

Approaches to valuing plant medicines: the economics of culture or the culture of economics?

This paper explores the scope and limitations of the economic valuation of biodiversity. How and why is this a useful exercise, but in what way is its usefulness constrained? These issues are discussed with particular reference to cultural values within the context of medicinal plants and phytopharmaceutical development. It is argued that whilst economic valuation is an extremely important and necessary aid to formulating conservation policy, it probably is not that informative about the motivations of people in their use of certain natural resources. This is especially true for the case of medicinal plants where belief systems about the causes of disease and therefore strategies for healing are important. The paper reviews studies which have attempted to put monetary values on medicinal plants and the option values of pharmaceuticals developed from plants, and discusses the applicability of the various approaches. The implications for conservation policy, and for likely collaboration between social and natural scientists is discussed.     

Primary or secondary? Versatile nitrilases in plant metabolism

The potential of plant nitrilases to convert indole-3-acetonitrile into the plant growth hormone indole-3-acetic acid has earned them the interim title of "key enzyme in auxin biosynthesis". Although not widely recognized, this view has changed considerably in the last few years. Recent work on plant nitrilases has shown them to be involved in the process of cyanide detoxification, in the catabolism of cyanogenic glycosides and presumably in the catabolism of glucosinolates. All plants possess at least one nitrilase that is homologous to the nitrilase 4 isoform of Arabidopsis thaliana. The general function of these nitrilases lies in the process of cyanide detoxification, in which they convert the intermediate detoxification product beta-cyanoalanine into asparagine, aspartic acid and ammonia. Cyanide is a metabolic by-product in biosynthesis of the plant hormone ethylene, but it may also be released from cyanogenic glycosides, which are present in a large number of plants. In Sorghum bicolor, an additional nitrilase isoform has been identified, which can directly use a catabolic intermediate of the cyanogenic glycoside dhurrin, thus enabling the plant to metabolize its cyanogenic glycoside without releasing cyanide. In the Brassicaceae, a family of nitrilases has evolved, the members of which are able to hydrolyze catabolic products of glucosinolates, the predominant secondary metabolites of these plants. Thus, the general theme of nitrilase function in plants is detoxification and nitrogen recycling, since the valuable nitrogen of the nitrile group is recovered in the useful metabolites asparagine or ammonia. Taken together, a picture emerges in which plant nitrilases have versatile functions in plant metabolism, whereas their importance for auxin biosynthesis seems to be minor.     

Does change in accessibility with conversion depend on both the substrate and pretreatment technology?

The accessibility of cellulase and xylanase enzymes to glucan and xylan, respectively, and its change with conversion were measured for pure Avicel glucan and poplar solids that had been pretreated by ammonia fiber expansion (AFEX), ammonia recycled percolation (ARP), dilute acid, and lime. Avicel and pretreated solids were digested to various degrees by cellulase together with β-glucosidase enzymes and then cleaned of residual protein via a biological method using Protease. Glucan accessibility was determined by purified CBHI (Cel7A) adsorption at 4 °C, and 4 and 24 h hydrolysis yields were determined for solids loading containing equal amounts of glucan (1.0% w/v) and lignin (1.0% w/v), in two separate sets of experiments. Consistent with our previous study and in contrast to some in the literature, little change in glucan accessibility was observed with conversion for Avicel, but glucan and xylan accessibility for real biomass varied with the type of pretreatment. For example, AFEX pretreated solids showed a negligible change in glucan accessibility for conversion up to 90%, although xylan accessibility seemed to decline first and then remained constant. On the other hand, a substantial decline in glucan and xylan accessibility with conversion was observed for lime pretreated poplar solids, as shown by initial hydrolysis rates. Yet, an increase in CBHI adsorption with conversion for lime pretreated poplar solids suggested the opposite trend, possibly due to increased lignin exposure and/or reduced effectiveness of adsorbed enzyme.     

Nitrogen transformations in boreal forest soils—does composition of plant secondary compounds give any explanations?

Two major groups of plant secondary compounds, phenolic compounds and terpenes, may according to current evidence mediate changes in soil C and N cycling, but their exact role and importance in boreal forest soils are largely unknown. In this review we discuss the occurrence of these compounds in forest plants and soils, the great challenges faced when their concentrations are measured, their possible effects in regulating soil C and N transformations and finally, we attempt to evaluate their role in connection with certain forest management practices. In laboratory experiments, volatile monoterpenes, in the concentrations found in the coniferous soil atmosphere, have been shown to inhibit net nitrogen mineralization and nitrification; they probably provide a C source to part of the soil microbial population but are toxic to another part. However, there is a large gap in our knowledge of the effects of higher terpenes on soil processes. According to results from laboratory experiments, an important group of phenolic compounds, condensed tannins, may also affect microbial processes related to soil C and N cycling; one mechanism is binding of proteins and certain other organic N-containing compounds. Field studies revealed interesting correlations between the occurrence of terpenes or phenolic compounds and C or net N mineralization in forest soils; in some cases these correlations point in the same direction as do the results from laboratory experiments, but not always. Different forest management practices may result in changes in both the quantity and quality of terpenes and phenolic compounds entering the soil. Possible effects of tree species composition, clear-cutting and removal of logging residue for bioenergy on plant secondary compound composition in soil are discussed in relation to changes observed in soil N transformations.     

How do herbivorous insects cope with noxious secondary plant compounds in their diet?

Herbivorous insects use a variety of physiological mechanisms to cope with noxious (i.e., unpalatable and/or toxic) compounds in their food plants. Here, I review what is known about this coping process, focusing on one species of caterpillar, the tobacco hornworm (Manduca sexta). Herbivorous insects possess both preingestive (i.e., chemosensory) and postingestive response mechanisms for detecting plant secondary compounds. Stimulation of either class of detection mechanism inhibits feeding rapidly by reducing biting rate and/or bite size. This aversive response is highly adaptive during encounters with secondary plant compounds that are toxic. The insect's dilemma is that many harmless or mildly toxic compounds also activate the aversive response. To overcome this dilemma, herbivorous insects employ at least three mechanisms for selectively deactivating their aversive response to relatively harmless secondary plant compounds: (1) the presence of carbohydrates can mask the unpalatable taste of some secondary plant compounds; (2) prolonged dietary exposure to some unpalatable secondary plant compounds can initiate long-term adaptation mechanisms in the peripheral and central gustatory system; and (3) dietary exposure to toxic compounds can induce production of P450 detoxication enzymes. Thus, herbivorous insects utilize an integrated suite of physiological mechanisms to detect potentially toxic compounds in foods, and then selectively adapt to those that do not pose a serious threat to their growth and survivorship.     

Old-field secondary succession in SE Spain: can fire divert it?

In the Mediterranean Basin, most cultivated areas were abandoned in the last century and are now in various stages of old-field succession. The aim of this work was to analyse the successional trajectories of these ecosystems, and to assess possible deviations in these pathways due to fire occurrence at high or low recurrence levels. Old-fields abandoned either about 50 or about 100 years ago were selected in SE Spain. Within the 50-year-old abandoned fields, plots were established which had been burned by 1, 2 and 3 fires in the last 25 years. Cover values of vascular species were sampled and then analysed by means of multivariate analysis. Euclidean distances between resulting communities were used as an indicator of the possible deviation from the unburned successional pathway. Our results pointed to the possibility that different successional pathways may exist depending on fire occurrence and recurrence. In the absence of fire, the vegetation is dominated by pioneer species, mainly Pinus. With the passage of time this vegetation will become dominated by later successional tree species (Quercus). However, when early-successional communities are affected by fire, the succession can be diverted. A single fire is enough to change Pinus forests into alternative stable states dominated by Rosmarinus officinalis shrub communities, where the colonisation of species in later successional stages is arrested. This deviation increases in high fire recurrence regimes where the vegetation changes to dwarf shrubs and herbs.     

Going with the wind – Adaptive dynamics of plant secondary meristems

The developmental plasticity of organisms is a natural consequence of adaptation. Classical approaches targeting developmental processes usually focus on genetics as the essential factor underlying phenotypic differences. However, such differences are often based on the inherent plasticity of developmental programs. Due to their dependence on environmental stimuli, plants represent ideal experimental systems in which to dissect the contribution of genetic and environmental variation to phenotypic plasticity. An evident example is the vast repertoire of growth forms observed in plant shoot systems. A fundamental factor underlying the broadness of this repertoire is the activity of secondary meristems, namely the axillary meristems that give rise to side shoots, and the cambium essential for stem thickening. Differential activities of both meristem types are crucial to the tremendous variation seen in higher plant architecture. In this review, we discuss the role of secondary meristems in the adaptation of plant growth forms, and the ways in which they integrate environmental input. In particular, we explore potential approaches for dissecting the degree to which this flexibility and its consequences for plant architecture is genetically predetermined and how much it represents an adaptive value.     

Does plant diversity increase top–down control of herbivorous insects in tropical forest?

Higher trophic level interactions are key mediators of ecosystem functioning in tropical forests. A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underlying such effects. The 'enemies hypothesis’ states that predators exert more effective top–down control of herbivorous insects with increasing plant diversity. Support for this hypothesis has been found in temperate forests and agroecosystems, but remains understudied in tropical forests. We compared incidence of attacks of different natural enemies using artificial caterpillars in a tropical forest landscape and investigated the role of plant community structure (i.e. species richness, composition and density), and the role of forest fragmentation (i.e. patch size, edge distance and canopy openness) on predation intensity. Plant community effects were tested with respect to three vegetation strata: trees, saplings and herbs. Observed predation was substantially due to ants. Predation rates increased with plant species richness for trees and herbs. Density of saplings, herb cover and herb species composition were important factors for predation. No significant patterns were found for fragmentation parameters, suggesting that forest fragmentation has not altered predation intensity. We conclude that in tropical forests, top–down control of herbivorous insects in the understory vegetation is affected by a combination of plant diversity, plant species composition and structural features of the plant community.     

Does the alternative pathway ameliorate chilling injury in sensitive plant tissues?

Free radical processes have been observed in senescence and several membrane-associated disorders of plants including chilling, freezing, and desiccation injuries. The mitochondria of plant tissues exposed to low temperatures, and other abiotic and biotic stresses, produce superoxide and/or hydrogen peroxide when electron transport through the cytochrome pathway is impaired due to the energy state of the cell or to stress-induced physical changes in the membrane components. The superoxide and/or hydrogen peroxide produced can diffuse throughout the cell causing peroxidation of membrane lipids which results in membrane disruption, increased permeability and metabolic disturbances, and eventually the visible symptoms of chilling injury. The alternative pathway of electron transport in the mitochondria, which is induced by low temperatures in some plant tissues, can mediate these degradative processes by reducing the level of superoxide generated by the mitochondria.     

Biotic resistance to plant invasion in grassland: Does seed predation increase with resident plant diversity?

Seed predation impacts heavily on plant populations and community composition in grasslands. In particular, generalist seed predators may contribute to biotic resistance, i.e. the ability of resident species in a community to reduce the success of non-indigenous plant invaders. However, little is known of predators’ preferences for seeds of indigenous or non-indigenous plant species or how seed predation varies across communities. We hypothesize that seed predation does not differ between indigenous and non-indigenous plant species and that seed predation is positively related to plant species diversity in the resident community. The seed removal of 36 indigenous and non-indigenous grassland species in seven extensively or intensively managed hay meadows across Switzerland covering a species-richness gradient of 18–50 plant species per unit area (c. 2 m²) was studied. In mid-summer 2011, c. 24,000 seeds were exposed to predators in Petri dishes filled with sterilized soil, and the proportions of seeds removed were determined after three days’ exposure. These proportions varied among species (9.2–62.5%) and hay meadows (17.8–48.6%). Seed removal was not related to seed size. Moreover, it did not differ between indigenous and non-indigenous species, suggesting that mainly generalist seed predators were active. However, seed predation was positively related to plant species richness across a gradient in the range of 18–38 species per unit area, representing common hay meadows in Switzerland. Our results suggest that generalist post-dispersal seed predation contributes to biotic resistance and may act as a filter to plant invasion by reducing the propagule pressure of non-local plant species.     

How does metal soil pollution change the plant mycobiome?

Microorganisms play a key role in plant adaptation to the environment. The aim of this study was to evaluate the effect of toxic metals present in the soil on the biodiversity of plant-related, endophytic mycobiota. The mycobiome of plants and soil from a Zn–Pb heap and a metal-free ruderal area were compared via Illumina sequencing of the ITS1 rDNA. The biodiversity of plants and fungi inhabiting mine dump substrate was lower than that of the metal free site. In the endosphere of Arabidopsis arenosa from the mine dump the number of endophytic fungal taxa was comparable to that in the reference population, but the community structure significantly differed. Agaricomycetes was the most notably limited class of fungi. The results of plant mycobiota evaluation from the field study were verified in terms of the role of toxic metals in plant endophytic fungi community assembly in a reconstruction experiment. The results presented in this study indicate that metal toxicity affects the structure of the plant mycobiota not by changing the pool of microorganisms available in the soil from which the fungal symbionts are recruited but most likely by altering plant and fungi behaviour and the organisms' preferences towards associating in symbiotic relationships.     

Does the Initial Floristic Composition model of succession really work?

Harvey & Holzman, in this issue of the Journal of Vegetation Science, describe divergent successional pathways after fire in Pinus muricata forest. Fourteen years after fire, the shrub Ceanothus thyrsiflorus had occupied flat parts of the area. They predicted it would continue to prevent succession to P. muricata forest, as in the initial floristic composition concepts of Clements and Egler.     

Does secondary structure determine tertiary structure in proteins?

Is highly approximate knowledge of a protein's backbone structure sufficient to successfully identify its family, superfamily, and tertiary fold? To explore this question, backbone dihedral angles were extracted from the known three‐dimensional structure of 2,439 proteins and mapped into 36 labeled, 60° × 60° bins, called mesostates. Using this coarse‐grained mapping, protein conformation can be approximated by a linear sequence of mesostates. These linear strings can then be aligned and assessed by conventional sequence‐comparison methods. We report that the mesostate sequence is sufficient to recognize a protein's family, superfamily, and fold with good fidelity. Proteins 2005. © 2005 Wiley‐Liss, Inc.     

The role of Quaternary environmental change in plant macroevolution: the exception or the rule?

The Quaternary has been described as an important time for genetic diversification and speciation. This is based on the premise that Quaternary climatic conditions fostered the isolation of populations and, in some instances, allopatric speciation. However, the 'Quaternary Ice-Age speciation model' rests on two key assumptions: (i) that biotic responses to climate change during the Quaternary were significantly different from those of other periods in Earth's history; and (ii) that the mechanisms of isolation during the Quaternary were sufficient in time and space for genetic diversification to foster speciation. These assumptions are addressed by examining the plant fossil record for the Quaternary (in detail) and for the past 410 Myr, which encompasses previous intervals of icehouse Earth. Our examination of the Quaternary record indicates that floristic responses to climate changes during the past 1.8 Myr were complex and that a distinction has to be made between those plants that were able to withstand the extremes of glacial conditions and those that could not. Generation times are also important as are different growth forms (e.g. herbaceous annuals and arborescent perennials), resulting in different responses in terms of genetic divergence rates during isolation. Because of these variations in the duration of isolation of populations and genomic diversification rates, no canonical statement about the predominant floristic response to climatic changes during the Quaternary (i.e. elevated rates of speciation or extinction, or stasis) is currently possible. This is especially true because of a sampling bias in terms of the fossil record of tree species over that of species with non-arborescent growth forms. Nevertheless, based on the available information, it appears that the dominant response of arborescent species during the Quaternary was extinction rather than speciation or stasis. By contrast, our examination of the fossil record of vascular plants for the past 410 Myr indicates that speciation rates often increased during long intervals of icehouse Earth (spanning up to 50 Myr). Therefore, longer periods of icehouse Earth than those occurring during the Quaternary may have isolated plant populations for sufficiently long periods of time to foster genomic diversification and allopatric speciation. Our results highlight the need for more detailed study of the fossil record in terms of finer temporal and spatial resolution than is currently available to examine the significance of intervals of icehouse Earth. It is equally clear that additional and detailed molecular studies of extant populations of Quaternary species are required in order to determine the extent to which these 'relic' species have genomically diversified across their current populations.     

Does plant height determine the freezing resistance in the páramo plants?

Neotropical ecosystems between treeline and snowline, called páramos, stretch along Andean ranges from Costa Rica to northern Peru. The páramo climate is characterized by regular night frosts occurring throughout the year. Páramo plants use two strategies to deal with freezing temperatures. They either avoid ice formation in the tissues or tolerate extracellular ice formation. We tested the microclimate hypothesis, which suggests that the freezing resistance of the páramo plants is determined by plant height, that is, that taller plants experience a milder microclimate and avoid freezing, whereas smaller plants are exposed to the more extreme thermal conditions near the ground and tolerate them. We measured the temperature at which ice formed inside the plants (the ‘exotherm’), and compared it with the temperature at which 50% damage to the tissue occurred (Lt50); a significant difference between the exotherm and Lt50 would indicate freezing tolerance whereas the absence of a difference would indicate avoidance by supercooling. We analysed the freezing resistance of 38 common Ecuadorian páramo species. We found no correlation between plant height and freezing resistance mechanism or injury temperature and reject the microclimate hypothesis. Tolerant plants reach higher altitudes than avoidant plants, but their altitudinal ranges largely overlap and the Lt50 does not differ between them. These results suggest that there is no qualitative difference between the two strategies to survive the páramo frosts. Shrub leaves were injured at significantly lower temperatures than other life forms, such as herbs, which may reflect leaf anatomical differences among the plants.     

Is subarctic forest advance able to keep pace with climate change?

Recent climate warming and scenarios for further warming have led to expectations of rapid movement of ecological boundaries. Here we focus on the circumarctic forest–tundra ecotone (FTE), which represents an important bioclimatic zone with feedbacks from forest advance and corresponding tundra disappearance (up to 50% loss predicted this century) driving widespread ecological and climatic changes. We address FTE advance and climate history relations over the 20th century, using FTE response data from 151 sites across the circumarctic area and site‐specific climate data. Specifically, we investigate spatial uniformity of FTE advance, statistical associations with 20th century climate trends, and whether advance rates match climate change velocities (CCVs). Study sites diverged into four regions (Eastern Canada; Central and Western Canada and Alaska; Siberia; and Western Eurasia) based on their climate history, although all were characterized by similar qualitative patterns of behaviour (with about half of the sites showing advancing behaviour). The main associations between climate trend variables and behaviour indicate the importance of precipitation rather than temperature for both qualitative and quantitative behaviours, and the importance of non‐growing season as well as growing season months. Poleward latitudinal advance rates differed significantly among regions, being smallest in Eastern Canada (~10 m/year) and largest in Western Eurasia (~100 m/year). These rates were 1–2 orders of magnitude smaller than expected if vegetation distribution remained in equilibrium with climate. The many biotic and abiotic factors influencing FTE behaviour make poleward advance rates matching predicted 21st century CCVs (~10³–10⁴ m/year) unlikely. The lack of empirical evidence for swift forest relocation and the discrepancy between CCV and FTE response contradict equilibrium model‐based assumptions and warrant caution when assessing global‐change‐related biotic and abiotic implications, including land–atmosphere feedbacks and carbon sequestration.