Growth rates of phytoplankton under fluctuating light

• 1 The effect of light fluctuations on the growth rates of four species of freshwater phytoplankton was investigated. Experimental light regimes included constant irradiance and fluctuations of a step function form, with equal proportion of high (maximum of 240 µmol photons m^‐2 s^‐1) and low light (minimum of 5 µmol photons m^‐2 s^‐1) (or dark) in a period. Fluctuations of 1, 8 and 24‐h periods were imposed over several average irradiances (25, 50, 100 and 120 µmol photons m^‐2 s^‐1).
• 2 Growth rate responses to fluctuations were species‐specific and depended on both the average irradiance and the period of fluctuations. Fluctuations at low average irradiances slightly increased growth rate of the diatom Nitzschia sp. and depressed growth of the cyanobacterium Phormidium luridum and the green alga Sphaerocystis schroeteri compared to a constant irradiance.
• 3 Fluctuations at higher average irradiance did not have a significant effect on the growth rates of Nitzschia sp. and Sphaerocystis schroeteri (fluctuations around saturating irradiances) and slightly increased the growth rates of the cyanobacteria Anabaena flos‐aquae and Phormidium luridum (when irradiance fluctuated between limiting and inhibiting levels).
• 4 In general, the effect of fluctuations tended to be greater when irradiance fluctuated between limiting and saturating or inhibiting levels of a species growth‐irradiance curve compared to fluctuations within a single region of the curve.
• 5 The growth rates of species under fluctuating light could not always be predicted from their growth‐irradiance curves obtained under constant irradiance. When fluctuations occur between limiting and saturating or inhibiting irradiances for the alga and when the period of fluctuations is long (greater than 8 h), steady‐state growth‐irradiance curves may be insufficient to predict growth rates adequately. Consequently, additional data on physiological acclimation, such as changes in photosynthetic parameters, may be required for predictions under non‐constant light supply in comparison to constant conditions.

     

An Incubator for the Simulation of a Fluctuating Light Climate in Studies of Planktonic Primary Productivity

A laboratory system for the quantification of phytoplankton photosynthesis under fluctuating light climate conditions is described. It consists of 2 temperature-controlled incubators with a variable light supply, an algal batch culture in incubation bottles with appropriate stirrers and a set of oxygen electrodes to monitor algal photosynthesis. By the rotation of special grey filters between the incubator and the light source, a regular up and down movement in the water column is simulated in up to 7 parallel bottles. Different ratios of euphotic depth to mixing depth and different velocities can be applied. Simultaneously, 8 bottles can be incubated under constant light. The system is demonstrated in experiments with Chlamydomonas sp. Further possibilities of application are proposed.     

Population and community responses of phytoplankton to fluctuating light

Light is a major resource in aquatic ecosystems and has a complex pattern of spatio-temporal variability, yet the effects of dynamic light regimes on communities of phytoplankton are largely unexplored. I examined whether and how fluctuating light supply affects the structure and dynamics of phytoplankton communities. The effect of light fluctuations was tested at two average irradiances: low, 25 μmol quanta m⁻² s⁻¹ and high, 100 μmol quanta m⁻² s⁻¹ in 2- and 18-species communities of freshwater phytoplankton. Species diversity, and abundances of individual species and higher taxa, depended significantly on both the absolute level and the degree of variability in light supply, while total density, total biomass, and species richness responded only to light level. In the two-species assemblage, fluctuations increased diversity at both low and high average irradiances and in the multispecies community fluctuations increased diversity at high irradiance but decreased diversity at low average irradiance. Species richness was higher under low average irradiance and was not affected by the presence or absence of fluctuations. Diatom abundance was increased by fluctuations, especially at low average irradiance, where they became the dominant group, while cyanobacteria and green algae dominated low constant light and all high light treatments. Within each taxonomic group, however, there was no uniform pattern in species responses to light fluctuations: both the magnitude and direction of response were species-specific. The temporal regime of light supply had a significant effect on the growth rates of individual species grown in monocultures. Species responses to the regime of light supply in monocultures qualitatively agreed with their abundances in the community experiments. The results indicate that the temporal regime of light supply may influence structure of phytoplankton communities by differentially affecting growth rates and mediating species competition. Received: 24 September 1997 / Accepted: 8 July 1998     

Reproductive interference in two ground-hopper species: testing hypotheses of coexistence in the field

Similar to resource competition, reproductive interference may hamper the coexistence of closely related species. Species that utilize similar signal channels during mate finding may face substantial fitness costs when they come into contact and demographic displacement of the inferior species (sexual exclusion) is a likely outcome of such interactions. The two ground‐hopper species Tetrix ceperoi and Tetrix subulata broadly overlap in their ranges and general habitat requirements, but rarely co‐occur on a local scale. Results from laboratory and field experiments suggest that this mosaic pattern of sympatry might be influenced by reproductive interference. Here, we examine the significance of sexual interactions for these species in the field and test hypotheses on mechanisms of coexistence. Our results show that heterospecific sexual interactions also occur under field conditions, but in contrast to the experiments T. ceperoi was not the inferior species. The number of male mating attempts of both species was strongly correlated with encounter frequencies. Males discriminated between the sexes but not between the species, suggesting an incomplete mate recognition system in both species. The analysis of microhabitat preferences and spatial distribution revealed that habitat partitioning is not a suitable mechanism of coexistence in this system. Instead, the costs of reproductive interference are substantially mitigated by different niche breadths leading to different degrees of aggregation. Despite a considerable niche overlap T. ceperoi displayed a stronger preference for bare ground and occurred more aggregated than T. subulata, which had a broader niche. These differences may reduce the frequencies of heterospecific encounters and interactions in the field. Our results demonstrate that coexistence in the presence of reproductive interference is comparable to resource competition, being strongly influenced by ecological traits of the involved species, such as niche breadth and dispersion pattern.     

Potential mechanisms of coexistence in closely related forbs

The stable coexistence of very similar species has perplexed ecologists for decades and has been central to the development of coexistence theory. According to modern coexistence theory, species can coexist stably (i.e. persist indefinitely with no long‐term density trends) as long as species' niche differences exceed competitive ability differences, even if these differences are very small. Recent studies have directly quantified niche and competitive ability differences in experimental communities at small spatial scales, but provide limited information about stable coexistence across spatial scales in heterogeneous natural communities. In this study, we use experimental and observational approaches to explore evidence for niche and competitive ability differences between two closely related, ecologically similar and widely coexisting annual forbs: Trachymene cyanopetala and T. ornata. We experimentally tested for stabilizing niche differences and competitive ability differences between these species by manipulating species' frequencies, under both well‐watered and water‐stressed conditions. We considered these experimental results in light of extensive field observations to explore evidence of niche segregation at a range of spatial scales. We found little evidence of intra‐specific stabilization or competitive ability differences in laboratory experiments while observational studies suggested niche segregation across pollinator assemblages and small‐scale microclimate heterogeneity. Though we did not quantify long‐term stabilization of coexisting populations of these species, results are consistent with expectations for stable coexistence of similar species via a spatial storage effect allowing niche differences to overcome even small (to absent) competitive ability differences.     

Leaf ontogenetic dependence of light acclimation in invasive and native subtropical trees of different successional status

In the Bonin Islands of the western Pacific where the light environment is characterized by high fluctuations due to frequent typhoon disturbance, we hypothesized that the invasive success of Bischofia javanica Blume (invasive tree, mid‐successional) may be attributable to a high acclimation capacity under fluctuating light availability. The physiological and morphological responses of B. javanica to both simulated canopy opening and closure were compared against three native species of different successional status: Trema orientalis Blume (pioneer), Schima mertensiana (Sieb. et Zucc.) Koidz (mid‐successional) and Elaeocarpus photiniaefolius Hook.et Arn (late‐successional). The results revealed significant species‐specific differences in the timing of physiological maturity and phenotypic plasticity in leaves developed under constant high and low light levels. For example, the photosynthetic capacity of T. orientalis reached a maximum in leaves that had just fully expanded when grown under constant high light (50% of full sun) whereas that of E. photiniaefolius leaves continued to increase until 50 d after full expansion. For leaves that had just reached full expansion, T. orientalis, having high photosynthetic plasticity between high and low light, exhibited low acclimation capacity under the changing light (from high to low or low to high light). In comparison with native species, B. javanica showed a higher degree of physiological and morphological acclimation following transfer to a new light condition in leaves of all age classes (i.e. before and after reaching full expansion). The high acclimation ability of B. javanica in response to changes in light availability may be a part of its pre‐adaptations for invasiveness in the fluctuating environment of the Bonin Islands.     

INTERACTIONS BETWEEN PULSED NUTRIENT SUPPLIES AND A PHOTOCYCLE AFFECT PHYTOPLANKTON COMPETITION FOR LIMITING NUTRIENTS IN LONG-TERM CULTURE1

The influence of periodic nutrient supplies and a photocycle on phytoplankton competition for limiting nutrients was examined using the diatoms Thalassiosira rotula Meunier (clone 411) and Chaetoceros sp. cf. vixvisibilis Schiller (clone 847). Chaetoceros sp. cf. vixvisibilis displaced T. rotula from ammonium-limited cultures under constant light irrespective of whether ammonium was supplied continuously, in 6 pulses.day^?1 or in a single daily pulse. In contrast, the species coexisted under the 14:10 h LD photocycle under either continuous or pulsed ammonium supplies with the relative abundance of C. sp. cf. vixvisibilis increasing as the interval between ammonium additions lengthened. Coexistence was not observed with either silicic acid or nitrate limitation. Chaetoceros sp. cf. vixvisibilis displaced T. rotula from both nitrate- and silicic acid-limited chemostat cultures and from semi-continuous cultures grown under the same photoperiod that produced coexistence with a daily pulse of ammonium. The presence of a photocycle was both necessary and sufficient to permit coexistence with ammonium limitation. Under continuous ammonium supply the photocycle may have induced a temporal separation of ammonium uptake between species, permitting sharing of the limiting nutrient and coexistence. In contrast, the species were shown to be in direct competition for the daily ammonium pulse. A competition model suggested that coexistence in this case arose from a balance between the species’ammonium uptake rates and their nitrogen demands for steady-state growth induced by the photocycle. The results indicate that variations in nutrient supply rates may contribute to the coexistence of phytoplankton species in the sea, but that the identity of the limiting nutrient and the influence of variations in other non-limiting resources play important roles in affecting the outcome of nutrient competition among planktonic algae.     

Allelopathy prevents competitive exclusion and promotes phytoplankton biodiversity

It has been hypothesized that allelopathy can prevent competitive exclusion and promote phytoplankton diversity in aquatic ecosystems, where numerous species coexist on a limited number of resources. However, experimental proof‐of‐principle is not available to support this hypothesis. Here we present the first experimental evidence to support this hypothesis by demonstrating that allelopathy promotes the coexistence of two phytoplankton species, Ankistrodesmus falcatus and Oscillatoria sp., that compete for a single limiting nutrient. By performing long‐term competition experiments in nitrate‐limited continuous cultures, and by describing the population dynamics using a mechanistic model, we demonstrate that when allelopathy comes into play, one of the following outcomes is possible depending on the relative initial abundances of the species: dominance of the stronger competitor for nitrate (the non‐allelopathic species), oscillatory coexistence, or dominance of the weaker competitor (the allelopathic species). Our model analysis revealed that sustained oscillatory coexistence of the two species would be a common outcome of this experiment. Our study confirms for the first time, based on laboratory experiments combined with mechanistic models, that allelopathy can alter the predicted outcome of inter‐specific competition in a nutrient‐limited environment and increase the potential for the coexistence of more species than resources, thereby contributing to the identification of endogenous mechanisms that explain the extreme diversity of phytoplankton communities.     

Niche breadth,rarity and ecological characteristics within a regional flora spanning large environmental gradients

Aim Species specialization, which plays a fundamental role in niche differentiation and species coexistence, is a key biological trait in relation to the responses of populations to changing environments. Species with a limited niche breadth are considered to experience a higher risk of extinction than generalist species. This work aims to measure the degree of specialization in the regional flora of the French Alps and test whether species specialization is related to species rarity and ecological characteristics. Location This study was conducted in the French Alps region, which encompasses a large elevational gradient over a relatively limited area (26,000 km²). Methods Specialization was estimated for approximately 1200 plant species found in the region. Given the inherent difficulty of pinpointing the critical environmental niche axes for each individual species, we used a co‐occurrence‐based index to estimate species niche breadths (specialization index). This comprehensive measurement included crucial undetermined limiting niche factors, acting on both local and regional scales, and related to both biotic and abiotic interactions. The specialization index for each species was then related to a selection of plant typologies such as Grime strategies and Raunkiaer life‐forms, and to two measurements of plant rarity, namely regional area of occupancy and local abundance. Results Specialist species were mainly found in specific and harsh environments such as wetlands, cold alpine habitats and dry heathlands. These species were usually geographically restricted but relatively dominant in their local communities. Although none of the selected traits were sufficient predictors of specialization, pure competitors were over‐represented amongst generalist species, whereas stress‐tolerant species tended to be more specialized. Main conclusions Our results suggest that co‐occurrence‐based indices of niche breadth are a satisfactory method for inferring plant specialization using large species samples across very heterogeneous environments. Our results are an empirical validation of the tolerance–dominance trade‐off and also provide interesting insights into the long‐standing question of which biological properties characterize species with narrow niche breadth that are potentially threatened by global changes in the environment.     

The influence of fluctuating light intensities on species composition and diversity of natural phytoplankton communities

The influence of fluctuating light intensities on phytoplankton composition and diversity was investigated for 49 days under semi-continuous culture conditions with sufficient nutrient supply, using phytoplankton assemblages from Lake Biwa, Japan. Light conditions were either periodically changed from high intensity (100 µmol photons m^-2 s^-1) to low intensity (20 µmol photons m^-2 s^-1) at intervals of 1, 3, 6 and 12 days, or fixed to constant intensities (permanent high and low light levels). All treatments additionally experienced a day:night cycle of 16:8 h. Phytoplankton abundance increased and reached a saturation level on day 19 of the treatment with permanent high light, but increased continuously until the end of the experiment (day 49) in the treatment with permanent low light intensity. In treatments with periodically changing light intensities, the phytoplankton abundance reached saturation levels between these dates. Under phytoplankton abundance saturation, chlorophytes predominated in the treatment with permanent high light, while either cyanophytes or diatoms were abundant under permanent low light intensity. Treatments with changing light supply had chlorophyte- and cyanobacteria-dominated replicates as well as replicates with balanced proportions of both. Furthermore, species diversity, measured by the Shannon index, was low in cultures under permanent light intensity, while slow fluctuating light at the scale of 3 -12 days resulted in an increased diversity index. These results indicate that species composition and diversity of the phytoplankton were affected by the periodically changing light regime in the order of days, and suggest that temporal changes in weather conditions are a major impediment to competitive exclusion of phytoplankton species in nature.     

Carbon allocation under light and nitrogen resource gradients in two model marine phytoplankton1

Marine phytoplankton have conserved elemental stoichiometry, but there can be significant deviations from this Redfield ratio. Moreover, phytoplankton allocate reduced carbon (C) to different biochemical pools based on nutritional status and light availability, adding complexity to this relationship. This allocation influences physiology, ecology, and biogeochemistry. Here, we present results on the physiological and biochemical properties of two evolutionarily distinct model marine phytoplankton, a diatom (cf. Staurosira sp. Ehrenberg) and a chlorophyte (Chlorella sp. M. Beijerinck) grown under light and nitrogen resource gradients to characterize how carbon is allocated under different energy and substrate conditions. We found that nitrogen (N)‐replete growth rate increased monotonically with light until it reached a threshold intensity (~200 μmol photons · m^?2 · s^?1). For Chlorella sp., the nitrogen quota (pg · μm^?3) was greatest below this threshold, beyond which it was reduced by the effect of N‐stress, while for Staurosira sp. there was no trend. Both species maintained constant maximum quantum yield of photosynthesis (mol C · mol photons^?1) over the range of light and N‐gradients studied (although each species used different photophysiological strategies). In both species, C:chl a (g · g^?1) increased as a function of light and N‐stress, while C:N (mol · mol^?1) and relative neutral lipid:C (rel. lipid · g^?1) were most strongly influenced by N‐stress above the threshold light intensity. These results demonstrated that the interaction of substrate (N‐availability) and energy gradients influenced C‐allocation, and that general patterns of biochemical responses may be conserved among phytoplankton; they provided a framework for predicting phytoplankton biochemical composition in ecological, biogeochemical, or biotechnological applications.     

Oxygen Productivity of Planktonic Algae under Fluctuating and Constant Light Conditions

Net oxygen productivity in cultures of Monoraphidium minutum, Cryptomonas sp. and Planktothrix agardhii exposed to fluctuating and constant light conditions was measured in a laboratory incubator. The fluctuating light climate simulated a linear up and down movement in a 2 m water column at 4 different ratios of euphotic depth to mixing depth. In addition, cultures were kept at a constant light climate simulating static incubation at 0, 0.5, 1 and 2 m depth and at the depth of the mean irradiance, respectively. Integral productivity in the simulated water column was lowest when algae were incubated at constant light in different depths, highest when the algae were incubated at constant mean photon flux density (PFD) and intermediate under fluctuating light. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Fluorescence in blue light (FLU) is involved in inactivation and localization of glutamyl‐tRNA reductase during light exposure

Fluorescent in blue light (FLU) is a negative regulator involved in dark repression of 5‐aminolevulinic acid (ALA) synthesis and interacts with glutamyl‐tRNA reductase (GluTR), the rate‐limiting enzyme of tetrapyrrole biosynthesis. In this study, we investigated FLU‘s regulatory function in light‐exposed FLU‐overexpressing (FLUOE) Arabidopsis lines and under fluctuating light intensities in wild‐type (WT) and flu seedlings. FLUOE lines suppress ALA synthesis in the light, resulting in reduced chlorophyll content, but more strongly in low and high light than in medium growth light. This situation indicates that FLU's impact on chlorophyll biosynthesis depends on light intensity. FLU overexpressors contain strongly increased amounts of mainly membrane‐associated GluTR. These findings correlate with FLU‐dependent localization of GluTR to plastidic membranes and concomitant inhibition, such that only the soluble GluTR fraction is active. The overaccumulation of membrane‐associated GluTR indicates that FLU binding enhances GluTR stability. Interestingly, under fluctuating light, the leaves of flu mutants contain less chlorophyll compared with WT and become necrotic. We propose that FLU is basically required for fine‐tuned ALA synthesis. FLU not only mediates dark repression of ALA synthesis, but functions also to control balanced ALA synthesis under variable light intensities to ensure the adequate supply of chlorophyll.     

Combined effects of light and nitrate supplies on the growth,photosynthesis and ultraviolet‐absorbing compounds in marine macroalga Gracilaria lemaneiformis (Rhodophyta), with special reference to the effects of solar ultraviolet radiation

It is well known that light and nutrients are essential to plants; however, there are few investigations in which these have been studied in combination on macroalgae, especially when solar ultraviolet radiation (UVR) is concerned. We cultured the red alga Gracilaria lemaneiformis (Bory) at different nitrate concentrations and light levels with or without UVR for 24 days. The results showed that nitrate supply markedly enhanced the growth and photosynthesis, increased the absorptivity of UV‐absorbing compounds (UVACs), and decreased photoinhibition in the presence of UVR. The thalli that received photosynthetically active radiation (PAR) treatment exhibited higher growth rates than those that received PAR + UVR at ambient or enhanced nitrate concentrations. However, under PAR + UVR treatment, the absorptivity of UVACs was higher than that of PAR and fluctuated with light levels. UVR was found to reduce the maximal net photosynthetic rate, apparent photosynthetic efficiency and light‐saturating irradiance while increasing the dark respiration rate, and inducing higher inhibition of growth and photosynthesis under high light versus under low light. Ultraviolet B significantly induced the synthesis of UVACs but led to higher inhibition on growth and photosynthesis than ultraviolet A.     

Acclimation of thermotolerant algae to light and temperature interaction1

Here, we explore the responses of photosynthesis and related cellular processes in the thermotolerant microalga Micractinium sp. acclimated to limiting and saturating irradiances combined with elevated temperatures, using a novel computer-controlled multi-sensor system. This system allows for the monitoring of online values of oxygen exchange during photosynthesis and respiration with high accuracy. Micractinium sp. cells showed maximum growth and net oxygen production rates under the optimal temperature of 25°C regardless of the light acclimation conditions. Our results show that the upper thermal threshold for Micractinium sp. photosynthesis and growth ranges between 35°C and 40°C. This microalga exhibited stable photosynthetic efficiency and effective non-photochemical quenching (NPQ) under saturating light, and was more susceptible to temperature change when acclimated to limiting light levels. These results demonstrate that the acclimation of thermotolerant microalgae to saturating light helps to enhance the thermal tolerance of PSII. This feature results from enhanced heat stability of PSII photochemistry and oxygen evolution.     

Dietary partitioning promotes the coexistence of planktivorous species on coral reefs

Theories involving niche diversification to explain high levels of tropical diversity propose that species are more likely to co‐occur if they partition at least one dimension of their ecological niche space. Yet, numerous species appear to have widely overlapping niches based upon broad categorizations of resource use or functional traits. In particular, the extent to which food partitioning contributes to species coexistence in hyperdiverse tropical ecosystems remains unresolved. Here, we use a molecular approach to investigate inter‐ and intraspecific dietary partitioning between two species of damselfish (Dascyllus flavicaudus, Chromis viridis) that commonly co‐occur in branching corals. Species‐level identification of their diverse zooplankton prey revealed significant differences in diet composition between species despite their seemingly similar feeding strategies. Dascyllus exhibited a more diverse diet than Chromis, whereas Chromis tended to select larger prey items. A large calanoid copepod, Labidocera sp., found in low density and higher in the water column during the day, explained more than 19% of the variation in dietary composition between Dascyllus and Chromis. Dascyllus did not significantly shift its diet in the presence of Chromis, which suggests intrinsic differences in feeding behaviour. Finally, prey composition significantly shifted during the ontogeny of both fish species. Our findings show that levels of dietary specialization among coral reef associated species have likely been underestimated, and they underscore the importance of characterizing trophic webs in tropical ecosystems at higher levels of taxonomic resolution. They also suggest that niche redundancy may not be as common as previously thought.     

Plastic and constant developmental traits contribute to adaptive differences in co-occurring Polygonum species

Adaptive differences among species are often thought to result from developmentally constant trait differences that enhance fitness in alternative environments. Species differences in patterns of individual phenotypic plasticity can also have ecological consequences. Indeed, functionally related constant and plastic traits may interact to determine the phenotype's adaptive value in particular conditions. We compared juvenile shade avoidance traits (height and its components, internode length and node number) across two field density treatments in Polygonumpersicaria and P. hydropiper, annual plant species that co‐occur in pastures comprised of a mosaic of plant densities. We used selection analyses to test trait contributions to fitness in alternative density treatments. Seedlings of both species expressed plasticity for internode elongation in response to density; P. persicaria plants increased internode length and consequently height significantly more in high density than did those of P. hydropiper. As predicted by the shade avoidance hypothesis, increased height was adaptive for both species in high density stands, so P. persicaria plants had higher fitness in this environment. By contrast, node numbers were relatively constant across density treatments in both species: P. hydropiper seedlings consistently produced more nodes than did those of P. persicaria. This constant trait difference contributed to P. hydropiper's greater relative fitness at low density, where more nodes and hence leaves enable plants to better exploit available light. Differences between species in these juvenile shade‐avoidance traits did not result from the evolutionary constraints of lack of heritable variation or costs of plasticity. We discuss how these interspecific trait differences may have been generated by divergent selective histories resulting from differences in herbivore resistance. These results illustrate how adaptive differences in both plastic and constantly expressed traits may jointly contribute to ecological distribution, including coexistence in patchy habitats.     

Trait adaptation promotes species coexistence in diverse predator and prey communities

Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time‐invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.     

Light interception in species with different functional groups coexisting in moorland plant communities

Competition for light is one of the most essential mechanisms affecting species composition. It has been suggested that similar light acquisition efficiency (Φ_mass, absorbed photon flux per unit aboveground mass) may contribute to species coexistence in multi-species communities. On the other hand, it is known that traits related with light acquisition vary among functional groups. We studied whether Φ_mass was similar among species with different functional groups coexisting in moorland communities. We conducted stratified clipping in midsummer when the stand biomass reached a maximum. Light partitioning among species was estimated using a model accounting for both direct and diffuse light. Evergreen species were found to have a significantly lower Φ_mass than deciduous species, which resulted from their lower absorbed photon flux per unit leaf area and lower specific leaf area. Shrubs had a smaller leaf mass fraction, but their Φ_mass was not lower than that of herbs because they had a higher leaf position due to the presence of wintering stems. Species with vertical leaves had a higher Φ_mass than those with horizontal leaves despite vertical leaves being a decided disadvantage in terms of light absorption. This higher Φ_mass was achieved by a greater leaf height in species with vertical leaves. Our results clearly demonstrate that light acquisition efficiency was different among the functional groups. However, the trend observed is not necessarily the same as that expected based on prior knowledge, suggesting that disadvantages in some traits for light acquisition efficiency are partly compensated for by other traits.     

THE INFLUENCE OF FLUCTUATING LIGHT ON DIVERSITY AND SPECIES NUMBER OF NUTRIENT‐LIMITED PHYTOPLANKTON1

The influence of fluctuating light on diversity and species number of a natural phytoplankton assemblage competing for nutrients was investigated for 48 days under semicontinuous culture conditions. Light conditions were either changed periodically from high (65 μmol photons·m^?2·s^?1) to low intensity (15 μmol photons·m^?2·s^?1) at intervals of 1, 3, 6, and 12 days or fixed at constant light conditions of intermediate intensity (40 μmol photons·m^?2·s^?1). Fluctuating light at intervals of 1–12 days significantly affected phytoplankton diversity. The development of phytoplankton communities differed in treatments with different light regimes. In treatments with long light intervals, species abundance oscillated with the light phases. Differences in the temporal development of phytoplankton communities resulted in hump‐shaped relations between the interval length of the light phases and both species number and diversity index and can be explained by the intermediate disturbance hypothesis. Fluctuating light tends to sustain phytoplankton diversity under nutrient limitation if the light regime changes in the order of several days. This indicates that temporal changes in weather regime are important in preventing competitive exclusion of phytoplankton species in nature.