Indirect Defense Responses to Herbivory in Grasses

The leaf trichome of tobacco (Nicotiana tabacum) represents a unique secretory structure in which the basal trichome cell is connected to the epidermis by numerous plasmodesmata (PD). Small fluorescent probes microinjected into the basal trichome cell moved apically into distal trichome cells but not into the subtending epidermal cell. In marked contrast, the same probes moved apically into trichome cells when injected into the epidermal cell. Noninvasive methods of dye loading, including ester loading into the apical secretory cell by trichome “capping” and by infiltration of caged fluorescein, produced the same result. In transgenic tobacco plants constitutively expressing photoactivatable green fluorescent protein (PAGFP), activation of PAGFP above the epidermal/trichome (e/t) boundary resulted in movement of protein apically into the distal trichome cells but not across the e/t boundary, while PAGFP activated in the epidermal cell moved apically across the e/t boundary. Experiments with apoplastic tracers also revealed the presence of a distinct apoplastic barrier to solute movement at the e/t interface. These data point to unidirectional transport of solutes through PD. PAGFP activated in individual cells equidistant between the basal cell and the apical cell moved bidirectionally from these cells, suggesting that mass flow was not the driving force for unidirectional transport. We found that unidirectional transport across the e/t boundary was not affected by virus infection or by addition of the actin inhibitor latrunculin but could be dissipated completely by addition of sodium azide. Collectively, our data suggest that active, unidirectional transport of molecules may occur through PD located at unique interfaces in the plant.Plasmodesmata (PD) are the functional units of the symplasm and occur at multiple interfaces between plant cells (Roberts, 2005; Maule, 2008). The basic structure of the PD pore reveals a plasma membrane (PM)-lined pore containing the desmotubule, an axial endoplasmic reticulum-derived structure that provides endomembrane continuity between cells (for review, see Maule, 2008). The space between the desmotubule and the PM is known as the cytoplasmic sleeve and is thought to be the main conduit for the movement of small solutes between cells (Erwee and Goodwin, 1983; Terry and Robards, 1987; Liarzi and Epel, 2005). PD have received considerable attention in recent years for their ability to traffic large macromolecules, including proteins and nucleic acids (Lucas and Lee, 2004; Maule, 2008). For many macromolecules, transport through the PD pore appears to occur by a selective mechanism that permits an exchange of macromolecules between specific cell types (Wu et al., 2002; Lucas and Lee, 2004; Zambryski, 2004).For small solutes, movement through the PD pore is thought to occur by diffusion and is strongly dependent on the hydrodynamic, or Stokes, radius of the permeant molecule in question (Liarzi and Epel, 2005). The effective channel for solute transport through PD is in the region of 3 nm (Erwee and Goodwin, 1983; Terry and Robards, 1987; Tucker, 1988), and molecules of less than 1 kD are thought to pass freely between cells (Erwee and Goodwin, 1983; Terry and Robards, 1987). However, a recalculation of the data presented by Terry and Robards (1987) by Fisher (1999) revealed that the effective diameter for transport through the cytoplasmic sleeve may be closer to 4 nm, potentially allowing significantly larger molecules to pass through the pore. Indeed, GFP (27 kD, Stokes radius of 2.8; Liarzi and Epel, 2005) may move diffusively between PD of some cell types, depending on tissue location and age (Oparka et al., 1999; Crawford and Zambryski, 2001; Wymer et al., 2001; Liarzi and Epel, 2005). Although the size exclusion limit of the PD pore was originally thought to be relatively constant (Erwee and Goodwin, 1983; Terry and Robards, 1987), it is now clear that “not all PD are equal,” in the sense that the size exclusion limit (SEL) of the pore may differ substantially during cell development (Duckett et al., 1994; Oparka et al., 1999; Crawford and Zambryski, 2000; Ruan et al., 2001) in response to environmental conditions (Epel and Erlanger, 1991; Cleland et al., 1994; Schulz, 1995) and the location of a given cell-cell interface in the plant (Erwee and Goodwin, 1983, 1985; Duckett et al., 1994; Ruan et al., 2001). It is also established that the SEL of the pore may respond to intracellular cues such as water relations (Oparka and Prior, 1992; Schulz, 1995), metabolic status (Tucker, 1993; Cleland et al., 1994; Wright and Oparka, 1997), cytosolic Ca²⁺ levels (Tucker, 1988; Tucker and Boss, 1996; Holdaway-Clarke et al., 2000), and hormones (Ormenese et al., 2006). The picture emerging is one in which the PD pore functions as a dynamic structure that alters its SEL in response to changing local conditions (Maule, 2008). Despite this, simple diffusion remains the most likely mechanism determining solute exchange between symplasmically coupled cells (Liarzi and Epel, 2005).A common feature in plant tissues is the presence of symplasmic “domains” within which PD share a common SEL (for review, see Roberts and Oparka, 2003). Examples of symplasmic domains occur at interfaces where cells of differing functions are connected by PD. In extreme cases, such as guard cells, PD are lost during development from the guard cell walls that connect the stomatal complex to the epidermis, effectively isolating the guard cells from the epidermal symplasm (Palevitz and Hepler, 1985). Commonly, however, cells of differing functions remain connected by PD, although these interfaces appear to be tightly regulated with respect to solute flow (Roberts and Oparka, 2003). An example of strict control occurs in the PD that connect the sieve element-companion cell complex with mesophyll cells (van Bel and van Rijen, 1994; Knoblauch and van Bel, 1998; Oparka and Turgeon, 1999). These PD appear to be closed during normal long-distance phloem functions (van Bel and van Rijen, 1994; Knoblauch and van Bel, 1998; Oparka and Turgeon, 1999) but may open in response to stress conditions (Schulz, 1995; Wright and Oparka, 1997). For those differing cell types that remain connected by PD, a major challenge is to retain cell autonomous functions in the presence of symplasmic connections.Recently, we made an extensive study of the interface that connects the basal leaf trichome cell of tobacco (Nicotiana tabacum) with the underlying epidermal cell (Faulkner et al., 2008). A simple freeze-fracture protocol allowed PD to be viewed in surface fractures of the cell wall and PM, and we were able to image all of the PD across the cell-cell interface of the epidermis/trichome (e/t) boundary. Our data showed that during wall development, simple primary PD were gradually replaced with secondary PD by a multiple twinning process that involved the de novo formation of PD pores. During trichome development, the number of PD at the e/t boundary increased 5-fold due to the insertion of new secondary PD pores. In mature trichomes, in excess of 2,000 PD were found in the cell wall spanning the e/t boundary. The ability to image large numbers of PD at this interface prompted us to explore more closely the function of PD at the e/t interface. Previously, trichomes have proved a useful experimental system for studying cell-cell transport through PD (Waigmann and Zambryski, 2000). Interestingly, trichome cells of Nicotiana clevelandii have been shown to be connected by PD with a large basal SEL of about 7 kD (Waigmann and Zambryski, 1995) and appear to differ in structure from mesophyll PD (Waigmann et al., 1997). However, the specific interface between the trichome basal cell and the epidermis has not been examined.To examine the function of PD at the e/t interface in more detail, we developed a number of different approaches for introducing fluorescent probes into the symplasm of the basal trichome cell and its supporting epidermal cell. These included microinjection, cell-specific dye loading, the use of caged fluorescein, and the generation of transgenic tobacco plants constitutively expressing photoactivatable GFP (PAGFP) in the cytosol. Our data show consistently that fluorescent probes introduced into the basal trichome cell are unable to cross the e/t boundary while probes introduced below this boundary are able to enter the trichome freely, providing strong evidence for unidirectional transport through PD at a specific cell-cell interface. In addition, we show that sodium azide, a metabolic inhibitor that alters PD SEL (Tucker, 1993), dissipates unidirectional solute flow into the trichome and allows molecules to exit the trichome. Our data are consistent with unidirectional solute movement into the basal trichome cell.     

The Myriad Plant Responses to Herbivores

Abstract Plant responses to herbivores are complex. Genes activated on herbivore attack are strongly correlated with the mode of herbivore feeding and the degree of tissue damage at the feeding site. Phloem-feeding whiteflies and aphids that produce little injury to plant foliage are perceived as pathogens and activate the salicylic acid (SA)-dependent and jasmonic acid (JA)/ethylene-dependent signaling pathways. Differential expression of plant genes in response to closely related insect species suggest that some elicitors generated by phloem-feeding insects are species-specific and are dependent on the herbivore's developmental stage. Other elicitors for defense-gene activation are likely to be more ubiquitous. Analogies to the pathogen-incompatible reactions are found. Chewing insects such as caterpillars and beetles and cell-content feeders such as mites and thrips cause more extensive tissue damage and activate wound-signaling pathways. Herbivore feeding is not equivalent to mechanical wounding. Wound responses are a part of the induced responses that accompany herbivore feeding. Herbivores induce direct defenses that interfere with herbivore feeding, growth and development, fecundity, and fertility. In addition, herbivores induce an array of volatiles that creates an indirect mechanism of defense. Volatile blends provide specific cues to attract herbivore parasites and predators to infested plants. The nature of the elicitors for volatile production is discussed.     

Signals Regulating Multiple Responses to Wounding and Herbivores

Damage inflicted by herbivore feeding necessitates multiple defense strategies in plants. The wound site must be sealed and defense responses mounted against the herbivore itself and against invading opportunistic pathogens. These defenses are controlled both in time and space by highly complex regulatory networks that themselves are modulated by interactions with other signaling pathways. In this review, we describe the signaling events that occur in individual wounded leaves, in systemic unwounded regions of the plant, and between the plant, and other organisms, and attempt to place these events in the context of a coordinated system. Key signals that are discussed include ion fluxes, active oxygen species, protein phosphorylation cascades, the plant hormones jasmonic acid, ethylene, abscisic acid and salicylic acid, peptide signals, glycans, volatile chemicals, and physical signals such as hydraulic and electrical signals. Themes that emerge after consideration of the published data are that glycans and peptide elicitors are likely primary triggers of wound-induced defense responses and that they function through the action of jasmonic acid, a central mediator of defense gene expression, whose effect is modulated by ethylene. In the field, wound signaling pathways are significantly impacted on by other stress response pathways, including pathogen responses that often operate through potentially antagonistic signals such as salicylic acid. However, gross generalisations are not possible because some wound and pathogen responses operate through common jasmonate- and ethylene-dependent pathways. Understanding the ways in which local and systemic wound signaling pathways are coordinated individually and in the context of the plants wider environment is a key challenge in the application of this science to crop-protection strategies.     

Multiple Effects of Introduced Mammalian Herbivores in a Temperate Forest

Introduced mammalian herbivores can significantly affect ecosystems. Here, I review evidence on effects of introduced mammalian herbivores in the temperate forest of the southern Andes. Available data suggest that introduced herbivores decrease the abundance of seedlings and saplings of dominant tree species in some forest types, which could impair forest regeneration. They also affect understory species composition. The mechanisms of the effects of introduced herbivores are complex, and include direct effects of browsing or trampling and more complex interactions such as indirect effects through other species. Some native mammalian and avian predators may benefit from increased food availability resulting from high densities of some introduced mammalian herbivores. In turn, enhanced populations of predators may have resulted in increased predation on native prey. Competition for resources and disease transmission have also been proposed as possible negative effects of introduced herbivores on native herbivores, but little evidence supports this claim. Little is known about effects on invertebrates.     

Recovering More than Tree Cover: Herbivores and Herbivory in a Restored Tropical Dry Forest

Intense and chronic disturbance may arrest natural succession, reduce environmental quality and lead to ecological interaction losses. Where natural succession does not occur, ecological restoration aims to accelerate this process. While plant establishment and diversity is promoted by restoration, few studies have evaluated the effect of restoration activities on ecological processes and animal diversity. This study assessed herbivory and lepidopteran diversity associated with two pioneer tree species growing in 4-year-old experimental restoration plots in a tropical dry forest at Sierra de Huautla, in Morelos, Mexico. The study was carried out during the rainy season of 2010 (July-October) in eleven 50 x 50 m plots in three different habitats: cattle-excluded, cattle-excluded with restoration plantings, and cattle grazing plots. At the beginning of the rainy season, 10 juveniles of Heliocarpus pallidus (Malvaceae) and Ipomoea pauciflora (Convolvulaceae) were selected in each plot (N = 110 trees). Herbivory was measured in 10 leaves per plant at the end of the rainy season. To evaluate richness and abundance of lepidopteran larvae, all plants were surveyed monthly. Herbivory was similar among habitats and I. pauciflora showed a higher percentage of herbivory. A total of 868 lepidopteran larvae from 65 morphospecies were recorded. The family with the highest number of morphospecies (9 sp.) was Geometridae, while the most abundant family was Saturnidae, with 427 individuals. Lepidopteran richness and abundance were significantly higher in H. pallidus than in I. pauciflora. Lepidopteran richness was significantly higher in the cattle-excluded plots, while abundance was significantly higher in the non-excluded plots. After four years of cattle exclusion and the establishment of plantings, lepidopteran richness increased 20 –fold in the excluded plots compared to the disturbed areas, whereas herbivory levels were equally high in both restored and disturbed sites. Restoration with plantings and exclusion of cattle and plantings was shown to be a successful strategy for attracting lepidopterans and cattle exclusion was the main factor explaining lepidopteran diversity.     

Complex Responses of Songbirds to Soil Acidification of Managed Beech Forests in Central Europe

Abstract Many forest ecosystems have been extensively degraded by acidifying pollutants over the last decades. There is also widespread concern about population decline of a number of songbirds due to acidification-driven calcium limitation. We hypothesize that the carrying capacity of forests decreases with reduced soil calcium content. We analyzed songbird assemblage/habitat relationships by territory mapping at 30 sites in a total of five managed beech forests. The sites harbored 38 songbird species. A canonical correspondence analysis showed that soil acidification, represented by soil calcium content, had a significant influence on the species assemblages of the 14 most frequent songbirds. The vertical complexity of the canopy additionally influenced the assemblages. Multiple regressions revealed that the density of the territories of song thrush (Turdus philomelos) and nuthatch (Sitta europaea) were positively correlated to calcium content. Blackbirds (Turdus merula) seemed to compensate for the negative effect of acidification by selecting breeding sites close to forest edges to forage in less acidified agricultural habitats. In contrast, the territorial density of robin (Erithacus rubecula), chaffinch (Phylloscopus collybita) and coal tit (Parus ater) increased with increasing soil acidification. Contrary to our hypothesis, the overall carrying capacity of songbird was not reduced in forests with acidified soils. However, the nesting success of song thrush and blackbird, as determined by the number of fledglings, was higher in the forest with the highest calcium content when compared to the forest with the lowest calcium content. We conclude that some species are severely affected by acidification, whereas others seem to have evolved successful strategies to meet their calcium demand even in acidified habitats. Electronic supplementary material: The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Gene Flow in European Beech (Fagus sylvatica L.)

Three relatively isolated stands were used to study gene flow in European beech (Fagus sylvatica L.) in Northern Germany. Nine allozyme loci (Got-B, Idh-A, Lap-A, Mdh-B, Mdh-C, Mnr-A, 6-pgdh-A, Pgi-B and Pgm-A) were utilized for multilocus-genotyping adult trees and seeds. Expected heterozygosity (He) ranged from 0.325 to 0.351 for the three stands. F(ST) revealed that there was small differentiation among stands (mean F(ST) = 0.013). The indirect estimates of gene flow (Nm) based on the mean F(ST) were high and the average Nm was 19.14. External gene flow by pollen ranged from 0.7 to 1.2% inferred from new alleles in seed samples. Moreover, paternity analysis was used to assess effective pollen dispersal by inferring paternity of offspring. The weighted mean distances of pollen dispersal for these three stands were 36.8 and 37.1 m based on simple exclusion procedure and most-likely method, respectively. Two of the trees in one stand had rare allozyme alleles (Lap-A1 and Idh-A4, respectively), which were used to directly measure pollen movement away from those trees. The frequency of the rare Lap and Idh alleles in seeds declines as the distance from the source tree increases. The weighted mean distance of pollen dispersal with rare allele Lap-A1 or Idh-A4 was 26.3 m.     

Moss Responses to Elevated CO2 and Variation in Hydrology in a Temperate Lowland Peatland

We studied the effects of elevated CO₂ (180–200 ppmv above ambient) on growth and chemistry of three moss species (Sphagnum palustre, S. recurvum and Polytrichum commune) in a lowland peatland in the Netherlands. Thereto, we conducted both a greenhouse experiment with both Sphagnum species and a field experiment with all three species using MiniFACE (Free Air CO₂ Enrichment) technology during 3 years. The greenhouse experiment showed that Sphagnum growth was stimulated by elevated CO₂ in the short term, but that in the longer term (≥1 year) growth was probably inhibited by low water tables and/or down-regulation of photosynthesis. In the field experiment, we did not find significant changes in moss abundance in response to elevated CO₂, although CO₂ enrichment appeared to reduce S. recurvum abundance. Both Sphagnum species showed stronger responses to spatial variation in hydrology than to increased atmospheric CO₂ concentrations. Polytrichum was insensitive to changes in hydrology. Apart from the confounding effects of hydrology, the relative lack of growth response of the moss species may also have been due to the relatively small increase in assimilated CO₂ as achieved by the experimentally added CO₂. We calculated that the added CO₂ contributed at most 32% to the carbon assimilation of the mosses, while our estimates based on stable C isotope data even suggest lower contributions for Sphagnum (24–27%). Chemical analyses of the mosses showed only small elevated CO₂ effects on living tissue N concentration and C/N ratio of the mosses, but the C/N ratio of Polytrichum was substantially lower than those of the Sphagnum species. Continuing expansion of Polytrichum at the expense of Sphagnum could reduce the C sink function of this lowland Sphagnum peatland, and similar ones elsewhere, as litter decomposition rates would probably be enhanced. Such a reduction in sink function would be driven mostly by increased atmospheric N deposition, water table regulation for agricultural purposes and land management to preserve the early successional stage (mowing, tree and shrub removal), since these anthropogenic factors will probably exert a greater control on competition between Polytrichum and Sphagnum than increased atmospheric CO₂ concentrations.     

Differential and Synergistic Functionality of Acylsugars in Suppressing Oviposition by Insect Herbivores

Acylsugars are secondary metabolites exuded from type IV glandular trichomes that provide broad-spectrum insect suppression for Solanum pennellii Correll, a wild relative of cultivated tomato. Acylsugars produced by different S. pennellii accessions vary by sugar moieties (glucose or sucrose) and fatty acid side chains (lengths and branching patterns). Our objective was to determine which acylsugar compositions more effectively suppressed oviposition of the whitefly Bemisia tabaci (Gennadius) (Middle East—Asia Minor 1 Group), tobacco thrips, Frankliniella fusca (Hinds), and western flower thrips, Frankliniella occidentalis (Pergande). We extracted and characterized acylsugars from four S. pennellii accessions with different compositions, as well as from an acylsugar-producing tomato breeding line. We also fractionated the acylsugars of one S. pennellii accession to examine the effects of its components. Effects of acylsugars on oviposition were evaluated by administering a range of doses to oviposition sites of adult whiteflies and thrips in non-choice and choice bioassays, respectively. The acylsugars from S. pennellii accessions and the tomato breeding line demonstrated differential functionality in their ability to alter the distribution of whitefly oviposition and suppress oviposition on acylsugar treated substrates. Tobacco thrips were sensitive to all compositions while western flower thrips and whiteflies were more sensitive to acylsugars from a subset of S. pennellii accessions. It follows that acylsugars could thus mediate plant-enemy interactions in such a way as to affect evolution of host specialization, resistance specificity, and potentially host differentiation or local adaptation. The acylsugars from S. pennellii LA1376 were separated by polarity into two fractions that differed sharply for their sugar moieties and fatty acid side chains. These fractions had different efficacies, with neither having activity approaching that of the original exudate. When these two fractions were recombined, the effect on both whiteflies and thrips exceeded the sum of the two fractions’ effects, and was similar to that of the original exudate. These results suggest that increasing diversity of components within a mixture may increase suppression through synergistic interactions. This study demonstrates the potential for composition-specific deployment of acylsugars for herbivore oviposition suppression, either through in planta production by tomato lines, or as biocides applied by a foliar spray.     

Land Use History Mediates Soil Biogeochemical Responses to Drought in Temperate Forest Ecosystems

Ecosystems - Terrestrial ecosystems are experiencing increasing frequency and intensity of droughts as a result of climate change. Despite a wealth of previous studies investigating soil responses...     

Vascular Plant Responses to Elevated CO2 in a Temperate Lowland Sphagnum Peatland

Vascular plant responses to experimental enrichment with atmospheric carbon dioxide (CO₂), using MINIFACE technology, were studied in a Dutch lowland peatland dominated by Sphagnum and Phragmites for 3 years. We hypothesized that vascular plant carbon would accumulate in this peatland in response to CO₂ enrichment owing to increased productivity of the predominant species and poorer quality (higher C/N ratios) and consequently lower decomposability of the leaf litter of these species. Carbon isotope signatures demonstrated that the extra 180 ppmv CO₂ in enriched plots had been incorporated into vegetation biomass accordingly. However, on the CO₂ sequestration side of the ecosystem carbon budget, there were neither any significant responses of total aboveground abundance of vascular plants, nor of any of the individual species. On the CO₂ release side of the carbon budget (decomposition pathway), litter quantity did not differ between ambient and CO₂ treatments, while the changes in litter quality (N and P concentration, C/N and C/P ratio) were marginal and inconsistent. It appeared therefore that the afterlife effects of significant CO₂-induced changes in green-leaf chemistry (lower N and P concentrations, higher C/N and C/P) were partly offset by greater resorption of mobile carbohydrates from green leaves during senescence in CO₂-enriched plants. The decomposability of leaf litters of three predominant species from ambient and CO₂-enriched plots, as measured in a laboratory litter respiration assay, showed no differences. The relatively short time period, environmental spatial heterogeneity and small plot sizes might explain part of the lack of CO₂ response. When our results are combined with those from other Sphagnum peatland studies, the common pattern emerges that the vascular vegetation in these ecosystems is genuinely resistant to CO₂-induced change. On decadal time-scales, water management and its effects on peatland hydrology, N deposition from anthropogenic sources and land management regimes that arrest the early successional phase (mowing, tree and shrub removal), may have a greater impact on the vascular plant species composition, carbon balance and functioning of lowland Sphagnum–Phragmites reedlands than increasing CO₂ concentrations in the atmosphere.     

Species Richness Pattern along Altitudinal Gradient in Central European Beech Forests

The unimodal species richness-altitude distribution pattern seems to be universal. To investigate the validity of this phenomenon in homogeneous substrate and vegetation conditions, we sampled beech-dominated forests in five volcanic mountain ranges in the Western Carpathians. European beech (Fagus sylvatica L.) formed monodominant closed-canopy stands at altitudes from 300 to 1,200 m. Along this gradient, the influence of beech on understory plant species richness was expected to be strong and uniform. The shape of the species richness-altitude relationship was analyzed for three datasets: herb layer, shrub layer, and both layers merged together. Contrary to prediction, the studied species richness-altitude relationship was inversely unimodal, with a minimum at intermediate altitudes. Quadratic regression models were statistically significant for all three datasets (P<0.001) and the explained variability ranged from 12 % to 20 %. The possible explanation for the observed pattern is twofold. In the central part of the altitudinal gradient, low species richness is due to strong competition by monodominant beech with accumulation of leaf litter and uptake soil resources, mainly water. This influence is somewhat released towards the margins of the gradient. Secondly, the species pool from the neighbouring communities increases species richness only in the lower parts of the altitudinal gradient.     

Subsurface CO<Subscript>2</Subscript> Dynamics in Temperate Beech and Spruce Forest Stands

Rates of soil respiration (CO₂ effluxes), subsurface pore gas CO₂/O₂ concentrations, soil temperature and soil water content were measured for 15 months in two temperate and contrasting Danish forest ecosystems: beech (Fagus sylvatica L.) and Norway spruce (Picea abies [L.] Karst.). Soil CO₂ effluxes showed a distinct seasonal trend in the range of 0.48–3.3 μmol CO₂ m⁻² s⁻¹ for beech and 0.50–2.92 μmol CO₂ m⁻² s⁻¹ for spruce and were well-correlated with near-surface soil temperatures. The soil organic C-stock (upper 1 m including the O-horizon) was higher in the spruce stand (184±23 Mg C ha⁻¹) compared to the beech stand (93±19 Mg C ha⁻¹) and resulted in a faster turnover time as calculated by mass/flux in soil beneath the beech stand (28 years) compared to spruce stand (60 years). Observed soil CO₂ concentrations and effluxes were simulated using a Fickian diffusion-reaction model based on vertical CO₂ production rates and soil diffusivity. Temporal trends were simulated on the basis of observed trends in the distribution of soil water, temperature, and live roots as well as temperature and water content sensitivity functions. These functions were established based on controlled laboratory incubation experiments. The model was successfully validated against observed soil CO₂ effluxes and concentrations and revealed that temporal trends generally could be linked to variations in subsurface CO₂ production rates and diffusion over time and with depths. However, periods with exceptionally high CO₂ effluxes (> 20 μmol CO₂ m⁻² s⁻¹) were noted in March 2000 in relation to drying after heavy rain and after the removal of snow from collars. Both cases were considered non-steady state and could not be simulated.     

Differential Recruitment of Camponotus femoratus (Fabricius) Ants in Response to Ant Garden Herbivory

Although several studies have shown that ants can recognize chemical cues from their host plants in ant-plant systems, it is poorly demonstrated in ant gardens (AGs). In this interaction, ant species constantly interact with various epiphyte species. Therefore, it is possible to expect a convergence of chemical signals released by plants that could be acting to ensure that ants are able to recognize and defend epiphyte species frequently associated with AGs. In this study, it was hypothesized that ants recognize and differentiate among chemical stimuli released by AG epiphytes and non-AG epiphytes. We experimentally simulated leaf herbivore damage on three epiphyte species restricted to AGs and a locally abundant understory herb, Piper hispidum, in order to quantify the number of recruited Camponotus femoratus (Fabricius) defenders. When exposed to the AG epiphytes Peperomia macrostachya and Codonanthe uleana leaves, it was observed that the recruitment of C. femoratus workers was, on average, respectively 556% and 246% higher than control. However, the number of ants recruited by the AG epiphyte Markea longiflora or by the non-AG plant did not differ from paper pieces. This indicated that ants could discern between chemicals released by different plants, suggesting that ants can select better plants. These results can be explained by evolutionary process acting on both ants’ capability in discerning plants’ chemical compounds (innate attraction) or by ants’ learning based on the epiphyte frequency in AGs (individual experience). To disentangle an innate behavior, a product of classical coevolutionary process, from an ant’s learned behavior, is a complicated but important subject to understand in the evolution of ant-plant mutualisms.     

Precipitation Pattern Regulates Soil Carbon Flux Responses to Nitrogen Addition in a Temperate Forest

Ecosystems - Changes in precipitation frequency and intensity are predicted to be more intense and frequent accompanying climate change and may have immediate or potentially prolonged effects on...