Orchids do not pay costs at emergence for prolonged dormancy

In plants, prolonged dormancy is often considered a response to resource depletion or environmental stress that comes at a fitness cost. However, apparent costs of dormancy could reflect the state in which plants entered dormancy, rather than effects of dormancy per se. We tested this hypothesis for a terrestrial orchid, Epipactis atrorubens, by analyzing differences in vital rates of dormant and emergent plants using generalized linear mixed models, applied to eight years of demographic data. Dormant E. atrorubens plants did not form one homogeneous stage class. Instead, the vital rates of dormant plants mirrored performance of plants in their life stage before dormancy. Plants emerging from dormancy were slightly (albeit only marginally statistically significantly) larger than plants transitioning from the matching aboveground stage class, especially for smaller and younger stage classes. Because small plants were most likely to go dormant, plants emerging from dormancy were also smaller than average, if one were to compare all previously dormant plants to all previously emergent plants. Therefore, misclassifying all dormant plants into a single stage class changes whether we view dormancy as intrinsically costly, in terms of future performance upon emergence. We suggest that prolonged dormancy may be a form of phenotypic plasticity in which plants distribute their performance and reproductive effort through time, rather than a simple stress response.     

Incorporation of plant carbon into the soil animal food web of an arable system

We used stable isotopes to examine the incorporation of plant carbon into the belowground food web of an agricultural system. Plots were established and planted with maize (Zea mays) in a rye field (Secale cereale) near G?ttingen (northern Germany) in May 1999. In October 1999, April 2000, and October 2000, meso- and macrofauna and maize and rye litter were collected in each plot and analyzed for 13C and 15N content. 15N signatures suggested that the soil animal species analyzed span three trophic levels with the trophic position of species varying little in time. The species investigated formed a continuum from primary to secondary decomposers to predators. On average, predator species differed from primary and secondary decomposers by 3.9 sigma15N suggesting that they fed on a mixed diet of both decomposer groups. The combined analysis of 13C and 15N signatures allowed us to identify links between prey and consumer species. In October 1999, shortly after maize residues had been incorporated into the plots, maize-born carbon was present in each of the animal species investigated, including top predators. The incorporation of maize carbon into the belowground food web increased during the following 12 months but the concentration of maize-born carbon never exceeded 50% in any of the species. Furthermore, the ranks of the incorporation of maize-born carbon of the species changed little. The results suggest that the belowground food web relies heavily on carbon originating from plant residues from before the recent two growing seasons. In most species the amount of maize-born carbon increased continuously; however, in some species it decreased during winter, suggesting that these species switched to a diet based more on C3 plants during winter, or predominantly metabolized carbon incorporated during the last growing season. The study documents that the combined analysis of 13C and 15N signatures in soil invertebrate species, after replacement of C3 by C4 plants, is a powerful tool to better understand the structure of the belowground food web and the flux of carbon through it.     

Carbohydrate storage and light requirements of tropical moist and dry forest tree species

In many plant communities, there is a negative interspecific correlation between relative growth rates and survival of juveniles. This negative correlation is most likely caused by a trade-off between carbon allocation to growth vs. allocation to defense and storage. Nonstructural carbohydrates (NSC) stored in stems allow plants to overcome periods of stress and should enhance survival. In order to assess how species differ in carbohydrate storage in relation to juvenile light requirements, growth, and survival, we quantified NSC concentrations and pool sizes in sapling stems of 85 woody species in moist semi-evergreen and dry deciduous tropical forests in the rainy season in Bolivia. Moist forest species averaged higher NSC concentrations than dry forest species. Carbohydrate concentrations and pool sizes decreased with the light requirements of juveniles of the species in the moist forest but not in the dry forest. Combined, these results suggest that storage is especially important for species that regenerate in persistently shady habitats, as in the understory of moist evergreen forests. For moist forest species, sapling survival rates increased with NSC concentrations and pool sizes while growth rates declined with the NSC concentrations and pool sizes. No relationships were found for dry forest species. Carbon allocation to storage contributes to the growth-survival trade-off through its positive effect on survival. And, a continuum in carbon storage strategies contributes to a continuum in light requirements among species. The link between storage and light requirements is especially strong in moist evergreen forest where species sort out along a light gradient, but disappears in dry deciduous forest where light is a less limiting resource and species sort out along drought and fire gradients.     

Annual life cycle of Chattonella spp., causative flagellates of noxious red tides in the Inland Sea of Japan

Effects of storage temperature on the dormancy and maturation of dormant cells of Chattonella spp. were investigated by the extinction dilution method using sediment samples collected in 1984 after the blooming in Harima-Nada. Seasonality of germinability in fresh sediments was also studied in 1985. A storage period of more than 4 months with a low temperature (11°C) was essential for the maturation to induce the germination capacity of dormant cells. The optimal temperature for germination (22°C) had no effects on the maturation. Using fresh Suo-Nada sediments, a marked seasonality of germinability was confirmed. It was weak from autumn to early winter, then strengthened gradually up to a high level, which was maintained between spring and early summer, and again decreased rapidly during summer. The annual life cycle of Chattonella spp., including vetetative and dormant phases, was summarized as follows: (1) vegetative cells in early summer originate from germination of dormant cells in sediments; (2) they form overwintering dormant cells during the summer season; (3) dormant cells spend a period of spontaneous dormancy until next spring; (4) the duration of post dormancy, an enforced one due to low temperatures, follows until early summer. The life cycle of Chattonella spp. is therefore well adapted to the temperature regime in temperate seas such as the Seto Inland Sea of Japan.     

多年生植物的芽休眠及调控机理研究进展

芽休眠是一种有益的生物学特性,能够使多年生植物避开不利的环境条件,使种群在不利环境条件下得以生存.综述了多年生植物芽休眠的机制及其调控的研究进展.芽休眠是一个受多个因素和多种基因综合调控的复杂生命现象,其形成和解除受到光照、温度、水分等外界环境因素的影响和调控.在这个过程中,植物体内的激素、糖类、多胺和Ca2+等信号分子以及抗氧化酶类和能量代谢酶类等发生一系列的变化与之相适应.这些变化是在相应的基因表达调控下实现的,这些基因涉及到光、温响应基因,水分响应基因,能量代谢基因,胁迫响应基因,激素相关基因,以及MADS-box转录因子等,它们构成一个复杂的调控网络,共同控制着芽休眠的形成与解除.结合研究课题提出今后芽休眠机制的研究方向以及在农业上的应用前景.图1参67     

Differences in diet composition and foraging patterns between sexes of the Magellanic penguin (Spheniscus magellanicus) during the non-breeding period as revealed by δ13C and δ15N values in feathers and bone

In diving seabirds, sexual dimorphism in size often results in sex-related differences of foraging patterns. Previous research on Magellanic penguins, conducted during the breeding season, failed to reveal consistent differences between the sexes on foraging behavior, despite sexual dimorphism. In this paper, we tested the hypothesis that male and female Magellanic penguins differ in diet and foraging patterns during the non-breeding period when the constraints imposed by chick rearing activities vanish. We used stable isotope ratios of carbon and nitrogen in feather and bone to characterize the diet and foraging patterns of male and female penguins in the South Atlantic at the beginning of the 2009–2010 and 2010–2011 post-breeding seasons (feathers) and over several consecutive breeding and migratory seasons (bone). The mean δ¹³C and δ¹⁵N values of feathers showed no differences between the sexes in any of the three regions considered or in the diet composition between the sexes from identical breeding regions; however, Bayesian ellipses showed a higher isotopic niche width in males at the beginning of the post-breeding season. Stable isotope ratios in bone revealed the enrichment of males with δ¹³C compared with females across the three regions considered. Furthermore, the Bayesian ellipses were larger for males and encompassed those of females in two of the three regions analyzed. These results suggest a differential use of winter resources between the sexes, with males typically showing a larger diversity of foraging/migratory strategies. The results also show that dietary differences between male and female Magellanic penguins may occur once the constraints imposed by chick rearing activities cease at the beginning of the post-breeding season.     

Long-term aboveground and belowground consequences of red wood ant exclusion in boreal forest

Despite their ubiquity, the role of ants in driving ecosystem processes both aboveground and belowground has been seldom explored, except within the nest. During 1995 we established 16 ant exclusion plots of approximately 1.1 x 1.1 m, together with paired control plots, in the understory layer of a boreal forest ecosystem in northern Sweden that supports high densities of the mound-forming ant Formica aquilonia, a red wood ant species of the Formica rufa group. Aboveground and belowground measurements were then made on destructively sampled subplots in 2001 and 2008, i.e., 6 and 13 years after set-up. While ant exclusion had no effect on total understory plant biomass, it did greatly increase the relative contribution of herbaceous species, most likely through preventing ants from removing their seeds. This in turn led to higher quality resources entering the belowground subsystem, which in turn stimulated soil microbial biomass and activity and the rates of loss of mass and carbon (C) and nitrogen (N) from litter in litterbags placed in the plots. This was accompanied by losses of approximately 15% of N and C stored in the humus on a per area basis. Ant exclusion also had some effects on foliar stable isotope ratios for both C and N, most probably as a consequence of greater soil fertility. Further, exclusion of ants had multitrophic effects on a microbe-nematode soil food web with three consumer trophic levels and after six years promoted the bacterial-based relative to the fungal-based energy channel in this food web. Our results point to a major role of red wood ants in determining forest floor vegetation and thereby exerting wide-ranging effects on belowground properties and processes. Given that the boreal forest occupies 11% of the Earth's terrestrial surface and stores more C than any other forest biome, our results suggest that this role of ants could potentially be of widespread significance for biogeochemical nutrient cycling, soil nutrient capital, and sequestration of belowground carbon.     

Constraints on global fire activity vary across a resource gradient

We provide an empirical, global test of the varying constraints hypothesis, which predicts systematic heterogeneity in the relative importance of biomass resources to burn and atmospheric conditions suitable to burning (weather/climate) across a spatial gradient of long-term resource availability. Analyses were based on relationships between monthly global wildfire activity, soil moisture, and mid-tropospheric circulation data from 2001 to 2007, synthesized across a gradient of long-term averages in resources (net primary productivity), annual temperature, and terrestrial biome. We demonstrate support for the varying constraints hypothesis, showing that, while key biophysical factors must coincide for wildfires to occur, the relative influence of resources to burn and moisture/weather conditions on fire activity shows predictable spatial patterns. In areas where resources are always available for burning during the fire season, such as subtropical/tropical biomes with mid-high annual long-term net primary productivity, fuel moisture conditions exert their strongest constraint on fire activity. In areas where resources are more limiting or variable, such as deserts, xeric shrublands, or grasslands/savannas, fuel moisture has a diminished constraint on wildfire, and metrics indicating availability of burnable fuels produced during the antecedent wet growing seasons reflect a more pronounced constraint on wildfire. This macro-scaled evidence for spatially varying constraints provides a synthesis with studies performed at local and regional scales, enhances our understanding of fire as a global process, and indicates how sensitivity to future changes in temperature and precipitation may differ across the world.     

Effects of four decades of fire manipulation on woody vegetation structure in Savanna

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.     

Modelling plant resource allocation and growth partitioning in response to environmental heterogeneity

Plant morphological adjustment in response to spatial resource heterogeneity is an important factor that determines the outcomes of plant–plant and plant–environment interactions. In this study, a dynamic model of resource allocation and growth partitioning at the whole-plant level is presented. The aim is to suggest a mechanism by which plants are capable of modifying their resource allocation to favour the growth of their growing parts sited in resource-rich patches. In this model, an individual plant is treated as a population of relatively independent subunits competing for internal resources. The growth decision of individual shoot and root subunits depends on their local endogenous nutrient status. The allocation of nutrients to different shoot parts and root parts is determined by the structure of the vascular networks. No specific partitioning functions and driving coefficients are introduced in the model to coordinate resource allocation and growth partitioning at the whole-plant level. Vascular tissues acquire resources from the nutrient flow passing through them to grow and maintain their activities. The simulation results show that, based on simple rules of nutrient supply, transport and utilization, plants are able to integrate activities at the whole-plant level to allocate proportionally more growth to their growing parts in the most favourable positions.     

人工景观生态湖滨净化带植物的遴选

对水生植物处理生活污水的效果等展开了一些初步的研究。利用盆栽试验,比较了香蒲、美人蕉等8种水生植物的净化效果,所选的8种植物对污水均有不同程度的去除效果。挺水植物对污水的净化效果和耐污能力均优于沉水植物。利用水生植物处理生活污水,5 d的停留时间可以使污水达到较好的去除效果。挺水植物中美人蕉和水竹对污水的净化效果比香蒲和灯心草更好;沉水植物中伊乐藻的净化效果最好。植物对污水的净化效率有明显的季节差异,冬季污水净化效率低于夏秋两季,沉水植物在不同季节脱氮除磷效果有显著的差异。     

Simulating the dynamics of sulfur species and zinc in wetland sediments

In situ measurements comparing vertical SO₄²⁻ profiles in vegetated and non-vegetated sediments showed that SO₄²⁻ concentrations in vegetated sediments increased significantly at the beginning of the growing season and then gradually decreased during the rest of the growing season. Throughout the growing season, SO₄²⁻ concentrations remained higher in the vegetated sediments than in the sediments without plants. The higher SO₄²⁻ concentrations in the vegetated sediments indicate that oxygen release from roots and evapotranspiration-induced advection by plants play an important role in the dynamics of sulfur species in sediments. Since the total pool of solid-phase sulfide is relatively large compared to the mass of SO₄²⁻ in the sediments, the gradual decrease of SO₄²⁻ concentrations may result from limitation of the solid-phase sulfide that is in direct contact with or very close to the roots and rhizomes. This would mean that the main pool of solid-phase sulfide and associated trace metals are not affected by the oxygen release from roots, and the associated trace metals will not become bioavailable during the growing season.     

碟形沼泽湿地水中氮的动态变化及影响因素分析

通过对三江平原碟形湿地不同植被类型沼泽水体中氮含量的研究表明,漂筏苔草、毛果苔草和乌拉苔草沼泽水体中各形态氮含量具有明显的季节变化特征.总氮(TN)含量在5-10月份的植物生长季节表现为先增加后降低的趋势,不同植被类型沼泽水体TN含量明显不同.NH4+-N是无机氮的主要存在形态,含量比例在44.81%～98.89%之间.在整个生长季,乌拉苔草沼泽水NH4+-N含量呈先增加后降低的趋势,毛果苔草沼泽水NH4+-N含量在8月份具有峰值,漂筏苔草沼泽水NH4+-N含量较低,波动不大.硝态氮(NO3--N)在生长季中期含量较低,而亚硝态氮(NO2--N)含量较高;生长季末期NO3--N含量增高,而NO2--N含量降低.沼泽水体中不同形态氮浓度与植被类型、植被生长阶段密切相关,并且受到水温、降雨、蒸发、农田排水等因素的影响.     

植物对沼泽湿地生态系统N2O排放的影响

利用静态暗箱/气相色谱法连续3个生长季(2003-2005年)对三江平原小叶章草甸和毛果苔草沼泽N2O排放通量进行野外对比观测试验.结果表明,植物不同生长阶段对湿地生态系统N2O排放通量的影响不尽相同.植物的参与促进了湿地生态系统N2O的排放,2003-2005年生长季小叶章草甸土壤-植物系统N2O排放通量分别是土壤表观N2O排放通量的1.58倍、2.09倍和2.34倍,同期毛果苔草沼泽土壤-植物系统N2O排放通量分别是土壤表观N2O排放通量的1.86倍、1.50倍和1.33倍.3个生长季小叶章草甸N2O排放通量均大于毛果苔草沼泽,这主要是由土壤理化性质的空间变异性以及水文情况的差异造成的.     

Resolving structural uncertainty in natural resources management using POMDP approaches

In recent years there has been a growing focus on the uncertainties of natural resources management, and the importance of accounting for uncertainty in assessing management effectiveness. This paper focuses on uncertainty in resource management in terms of discrete-state Markov decision processes (MDP) under structural uncertainty and partial observability. It describes the treatment of structural uncertainty with approaches developed for partially observable resource systems. In particular, I show how value iteration for partially observable MDPs (POMDP) can be extended to structurally uncertain MDPs. A key difference between these process classes is that structurally uncertain MDPs require the tracking of system state as well as a probability structure for the structure uncertainty, whereas with POMDPs require only a probability structure for the observation uncertainty. The added complexity of the optimization problem under structural uncertainty is compensated by reduced dimensionality in the search for optimal strategy. A solution algorithm for structurally uncertain processes is outlined for a simple example in conservation biology. By building on the conceptual framework developed for POMDPs, natural resource analysts and decision makers who confront structural uncertainties in natural resources can take advantage of the rapid growth in POMDP methods and approaches, and thereby produce better conservation strategies over a larger class of resource problems.     

Population and individual consequences of breeding resource availability in the European bitterling (Rhodeus amarus)

Resource availability may affect both individual fitness and population demography and the effects can interact. We used two experiments to test how breeding resource abundance and its spatial distribution, combined with female abundance, affected male reproductive behavior, population spawning rate, and embryo development and recruitment in the European bitterling (Rhodeus amarus), a small cyprinid fish that lays its eggs in living unionid mussels. In the first experiment, we found that at the population level the abundance of breeding resources (freshwater mussels) was more important for bitterling recruitment than resource spatial distribution (clumped or regular). In contrast at the individual level, (variability in reproductive success) the spatial distribution of resources was more important, but only when resource abundance was not limiting. Territorial males obtained almost exclusive access to fertilizations when resources were abundant and distributed regularly, but were unable to defend large clusters of resources (when rival abundance was always high) and abandoned territoriality. Surprisingly, territorial males remained aggressive and successfully defended their territories when resources were grouped into a single cluster, but at a low abundance. In the second experiment, more rapid embryo development and larger juvenile body size at the end of the growing season was detected at high resource abundance and low female abundance, indicating that early hatched juveniles survived better and hence investment in offspring production early in the season yields a higher fitness pay-offs. The abundance of females in spawning condition was the best overall predictor of the intensity of male reproductive behavior in both experiments.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers

The timing of life history traits is central to lifetime fitness and nowhere is this more evident or well studied as in the phenology of flowering in governing plant reproductive success. Recent changes in the timing of environmental events attributable to climate change, such as the date of snowmelt at high altitudes, which initiates the growing season, have had important repercussions for some common perennial herbaceous wildflower species. The phenology of flowering at the Rocky Mountain Biological Laboratory (Colorado, USA) is strongly influenced by date of snowmelt, which makes this site ideal for examining phenological responses to climate change. Flower buds of Delphinium barbeyi, Erigeron speciosus, and Helianthella quinquenervis are sensitive to frost, and the earlier beginning of the growing season in recent years has exposed them to more frequent mid-June frost kills. From 1992 to 1998, on average 36.1% of Helianthella buds were frosted, but for 1999-2006 the mean is 73.9%; in only one year since 1998 have plants escaped all frost damage. For all three of these perennial species, there is a significant relationship between the date of snowmelt and the abundance of flowering that summer. Greater snowpack results in later snowmelt, later beginning of the growing season, and less frost mortality of buds. Microhabitat differences in snow accumulation, snowmelt patterns, and cold air drainage during frost events can be significant; an elevation difference of only 12 m between two plots resulted in a temperature difference of almost 2 degrees C in 2006 and a difference of 37% in frost damage to buds. The loss of flowers and therefore seeds can reduce recruitment in these plant populations, and affect pollinators, herbivores, and seed predators that previously relied on them. Other plant species in this environment are similarly susceptible to frost damage so the negative effects for recruitment and for consumers dependent on flowers and seeds could be widespread. These findings point out the paradox of increased frost damage in the face of global warming, provide important insights into the adaptive significance of phenology, and have general implications for flowering plants throughout the region and anywhere climate change is having similar impacts.     

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.