Seasonal Changes In Light Availability Research Articles

Photoperiodic dependent regulation of photosynthesis in the polar diatom Fragilariopsis cylindrus

Introduction: Polar microalgae are exposed to dramatic seasonal changes in light availability, from continuous summer days to winter nights with rapid changes of the daylength in spring and fall. Under this challenging light climate, large diatoms spring blooms occur at the bottom sea-ice and underneath the icepack, accounting for a significant proportion of the annual marine primary production in the Arctic Ocean. The on-going earlier melt down of the snow and ice covers result in a stronger light penetration and consequent increase in irradiance at the bottom of the sea ice leading to earlier seasonal sea-ice diatom blooms under shorter daylengths. Therefore, elucidating the response of polar diatoms to different photoperiods will help to better understand the consequences of the changing arctic climate on their photosynthetic productivity.Methods: In this study, we characterized the response of F. cylindrus, a model polar diatom, across five different photoperiods with similar light and temperature conditions (30 μmol photons m-2 s-1 and 0°C respectively).Results: We report different photoacclimative strategies under shorter and longer daylengths, with the special case of prolonged darkness (mimicking winter polar night). We also observed a repeated daily regulation of the photochemistry and photoprotection parameters when cells were exposed to a light:darkness alternation, despite the constant and optimal light intensity during the light periods.Discussion: Our results highlight the ability of F. cylindrus to grow efficiently under a wide range of daylengths, finely adjusting the balance between photochemistry and photoprotection to make the best use of the available light, supporting sustained production and growth despite low light and temperature.

Trade-offs Between Light and Nutrient Availability Across Gradients of Dissolved Organic Carbon Lead to Spatially and Temporally Variable Responses of Lake Phytoplankton Biomass to Browning

Northern lakes are experiencing widespread increases in dissolved organic carbon (DOC) that are likely to lead to changes in pelagic phytoplankton biomass. Pelagic phytoplankton biomass responds to trade-offs between light and nutrient availability. However, the influence of DOC light absorbing properties and carbon–nutrient stoichiometry on phytoplankton biomass across seasonal or spatial gradients has not been assessed. Here, we analyzed data from almost 5000 lakes to examine how the carbon–phytoplankton biomass relationship is influenced by seasonal changes in light availability, DOC light absorbing properties (carbon-specific visual absorbance, SVA420), and DOC–nutrient [total nitrogen (TN) and total phosphorus (TP)] stoichiometry, using TOC as a proxy for DOC. We found evidence for trade-offs between light and nutrient availability in the relationship between DOC and phytoplankton biomass [chlorophyll (chl)-a], with the shape of the relationship varying with season. A clear unimodal relationship was found only in the fall, particularly in the subsets of lakes with the highest TOC:TP. Observed trends of increasing TOC:TP and decreasing TOC:TN suggest that the effects of future browning will be contingent on future changes in carbon–nutrient stoichiometry. If browning continues, phytoplankton biomass will likely increase in most northern lakes, with increases of up to 76% for a 1.7 mg L−1 increase in DOC expected in subarctic regions, where DOC, SVA420, DOC:TN, and DOC:TP are all low. In boreal regions with higher DOC and higher SVA420, and thus lower light availability, lakes may experience only moderate increases or even decreases in phytoplankton biomass with future browning.

Seasonal changes in light availability modify the temperature dependence of secondary production in an Arctic stream.

Light and temperature are key drivers of ecosystem productivity, but synchrony of their annual cycles typically obscures their relative influence. The coupling of annual light-temperature regimes also drives complementary seasonal cycles of energy supply (primary production) and demand (metabolism), perhaps promoting temporal stability in carbon (C) storage and food web production that may be difficult to discern in most ecosystems. Spring-fed streams in the Arctic are subject to extreme annual fluctuations in light availability but have relatively stable water temperatures, which allows assessment of the independent effects of light and temperature. We used the unusual annual light and temperature regimes of Ivishak Spring, Alaska, USA (latitude 69°N, annual water temperature range ~4-7°C) to test predictions about the effect of light availability on consumer productivity with minimally confounding effects of temperature. We predicted that (1) annual patterns of secondary production would follow patterns of primary production, rather than temperature, due to organic C limitation during winter darkness when photosynthesis is effectively halted, (2) C limitation would propagate from primary producers upward through several trophic levels, (3) the lack of temperature dependence during winter darkness would be expressed as anomalous Arrhenius plots of growth rates indicating decoupled production-temperature relationships, and (4) consumer diets would reflect C limitation during winter. As predicted, we found (1) lowest production by macroinvertebrates and Salvelinus malma (Dolly Varden char) at the lowest light levels rather than the lowest temperatures, (2) apparent winter C limitation propagated upward through three trophic levels, (3) anomalous Arrhenius plots indicating lack of temperature dependence of consumer growth rates during winter, and (4) lowest consumption of diatoms (by macroinvertebrates) and invertebrate prey (by S.malma) during winter. Together, these results indicate that light drives annual patterns of animal production in Ivishak Spring, with stable annual temperatures likely exacerbating C limitation of ectotherm metabolism during winter. The timing and severity of winter C limitation in this unusual Arctic-spring food web highlight a fundamental role for light-temperature synchrony in matching energy supply with demand in most other ecosystem types, thereby conferring a measure of stability in the metabolism of their food webs over annual time scales.

Amazon forest structure generates diurnal and seasonal variability in light utilization

Abstract. The complex three-dimensional (3-D) structure of tropical forests generates a diversity of light environments for canopy and understory trees. Understanding diurnal and seasonal changes in light availability is critical for interpreting measurements of net ecosystem exchange and improving ecosystem models. Here, we used the Discrete Anisotropic Radiative Transfer (DART) model to simulate leaf absorption of photosynthetically active radiation (lAPAR) for an Amazon forest. The 3-D model scene was developed from airborne lidar data, and local measurements of leaf reflectance, aerosols, and PAR were used to model lAPAR under direct and diffuse illumination conditions. Simulated lAPAR under clear-sky and cloudy conditions was corrected for light saturation effects to estimate light utilization, the fraction of lAPAR available for photosynthesis. Although the fraction of incoming PAR absorbed by leaves was consistent throughout the year (0.80–0.82), light utilization varied seasonally (0.67–0.74), with minimum values during the Amazon dry season. Shadowing and light saturation effects moderated potential gains in forest productivity from increasing PAR during dry-season months when the diffuse fraction from clouds and aerosols was low. Comparisons between DART and other models highlighted the role of 3-D forest structure to account for seasonal changes in light utilization. Our findings highlight how directional illumination and forest 3-D structure combine to influence diurnal and seasonal variability in light utilization, independent of further changes in leaf area, leaf age, or environmental controls on canopy photosynthesis. Changing illumination geometry constitutes an alternative biophysical explanation for observed seasonality in Amazon forest productivity without changes in canopy phenology.

Seasonal changes in light availability modify the temperature dependence of ecosystem metabolism in an arctic stream

Light and temperature are key ecosystem drivers, but their synchronous annual cycles typically confound partitioning of their relative influence. Arctic spring‐streams, subject to extreme annual fluctuations in light but stable water temperatures, provide a rare contrast that allows the parsing of their independent effects. Over 30 months, we assessed the effects of light and temperature on ecosystem metabolism and nutrient uptake in Ivishak Spring, Alaska, USA. (latitude 69° N, water temperature range ∼4°–7°C) using open‐channel methods and short‐term NH4+‐N, NO3−‐N, and P additions, respectively. We predicted that rates of ecosystem respiration (ER) would mirror seasonal patterns of gross primary production (GPP), rather than temperature, due to relatively constant rates of metabolic demand year‐round, resulting in carbon limitation during winter (October–March) when photosynthesis effectively ceases. Because patterns of nutrient uptake and GPP are often coupled due to assimilatory demand, we also predicted that extreme annual cycles of light would result in equally extreme cycles of nutrient uptake, with demand being relaxed during winter.In accordance with our prediction, we found that ER scaled linearly with GPP. Peak summer rates of GPP (>4.0 g C·m−2·d−1) and ER (>5.0 g C·m−2·d−1) were surprisingly high, being comparable to those of productive streams at temperate latitudes. Winter rates (GPP ∼0.0, ER <1.0 g C·m−2·d−1) were low, however, and Arrhenius plots showed clear deviations from theoretical temperature dependence of GPP and ER during winter when other factors assumed primacy. For GPP, this factor was undoubtedly light availability, but for ER, carbon limitation is implicated due to low GPP. Significant nutrient uptake occurred only for NH4+‐N, indicating N limitation, and rates of uptake were also synchronous with cycles of light availability. Consequently, light, rather than temperature, was the major driver of annual patterns of ER and nutrient cycles in this arctic ecosystem. Synchronous light and temperature cycles are pervasive among ecosystems. The winter onset and severity of energy limitation we document highlights the importance of this synchrony and how the confounding of light and temperature obscures details of mechanisms by which these fundamental drivers affect ecosystem processes.

Seasonal influences on carbohydrate metabolism in the CAM bromeliad Aechmea 'Maya': consequences for carbohydrate partitioning and growth.

Photosynthetic plasticity in response to a range of environmental factors that include [CO(2)], water availability, light intensity and temperature, is ubiquitous among plants with crassulacean acid metabolism (CAM). The present study examined how seasonal changes in light availability, as experienced by greenhouse CAM crops in northern latitude regions, influence diel carboxylation patterns and impact on carbon gain and seasonal accumulation of biomass. In the CAM bromeliad Aechmea 'Maya' integrated measurements of leaf gas exchange, diel metabolite dynamics (e.g. malate, soluble sugars and starch) and biomass accumulation were made four times a year, i.e. in winter, spring, summer and autumn. During the brighter seasons (spring and summer) daytime Phases II and IV were dominated by C(4) carboxylation, whilst the higher diurnal uptake in the autumn and winter was characterized by equal contributions of both Rubisco and PEPC. As a consequence, net CO(2) uptake showed a significant depression at the end of the day in the darker months when supplementary illumination was turned off. Remarkable seasonal consistency was found in the amount of storage reserves available for nocturnal carboxylation, a consequence of predominantly daytime export of carbohydrate in spring and summer whilst nocturnal export was the major sink for carbohydrate in autumn and winter. Throughout the different seasons Aechmea 'Maya' showed considerable plasticity in the timing and magnitude of C(3) and C(4) carboxylation processes over the diel cycle. Under low PPFD (i.e. winter and autumn) it appears that there was a constraint on the amount of carbohydrate exported during the day in order to maintain a consistent pool of transient carbohydrate reserves. This gave remarkable seasonal consistency in the amount of storage reserves available at night, thereby optimizing biomass gain throughout the year. The data have important practical consequences for horticultural productivity of CAM plants and suggest a scenario for reconciling carbohydrate partitioning between competing sinks of nocturnal acidification and export for growth.

Flowering and fruiting phenologies of seasonal and aseasonal neotropical forests: the role of annual changes in irradiance

The seasonality of both rainfall and solar irradiance might influence the evolution of flowering and fruiting in tropical forests. In seasonally dry forests, to the degree that soil moisture limits plant productivity, community-wide peaks in reproduction are expected during the rainy season, with seedfall and germination timed to allow seedlings to become well established while soil moisture is available. Where soil moisture is never seasonally limiting, seasonal changes in light availability caused by periods of cloudiness or seasonally low zenithal sun angles should favour reproduction during seasons when irradiance levels are high. To evaluate these predictions, we documented the timing of flower and fruit fall for 10 and 15 y at El Verde, Puerto Rico, and Barro Colorado Island (BCI), Panama. At El Verde, rainfall is abundant year-round and solar declination largely determines seasonal variation in irradiance. At BCI, rainfall is abundant throughout the 8-mo wet season while drought develops and average solar irradiance increases by 40–50% over the 4-mo dry season. Seasonal variation in the number of species flowering and fruiting at both sites was generally consistent with the hypothesis that seasonal variation in irradiance limits the evolution of reproductive phenologies. Community-level metrics provided no evidence for a similar role for moisture availability at BCI. Seasonal variation in irradiance also strongly influenced seed development times at both sites. Thus, community-wide phenologies reveal a strong signature of seasonal changes in irradiance, even in those forests that exhibit some degree of seasonality in rainfall.

Do seasonal changes in light availability influence the inverse leafing phenology of the neotropical dry forest understory shrub Bonellia nervosa (Theophrastaceae)?

In tropical dry forests most plants are deciduous during the dry season and flush leaves with the onset of the rains. In Costa Rica, the only species displaying the opposite pattern is Bonellia nervosa. To determine if seasonal changes in light availability are associated with the leaf and reproductive phenology of this species, we monitored leaf production, survival, and life span, as well as flower and fruit production from April 2000 to October 2001 in Santa Rosa National Park. Leaf flushing and flower bud production took place shortly after the autumnal equinox when day length starts to decrease. Leaves began expansion at the end of the wet season, and plants reached 70 % of their maximum leaf area at the beginning of the dry season, maintaining their foliage throughout the entire dry period. Leaf shedding occurred gradually during the first three months of the wet season. Leaf flushing and shedding showed high synchrony, with leaf numbers being related to light availability. Maximum leaf production coincided with peaks in radiation during the middle of the dry season. Decreasing day length induces highly synchronous flower bud emergence in dry forest species, but this is the first study indicating induction of leaf flushing by declining day length.

Influence of canopy tree phenology on understorey populations of Fagus crenata

Abstract:Question: Is light available for subcanopy individuals of Fagus crenata spatiotemporally heterogeneous across patches with closed canopies of different foliage phenologies and gaps? Is local abundance of Sasa influenced by the composition of the canopy layer? If so, does the Sasa layer also affect the amount of light available to small F. crenata saplings? Is variation in F. crenata population structure consistent with the hypothesis that light is important?Location: Mt. Kurikoma, Japan 780 m a.s.l.Methods: Population structure of subcanopy individuals of Fagus crenata and importance of Sasa were examined in five patch types. The patch‐types were Fc (F. crenata only in the crown), Qm (Quercus mongolica var. grosseserrata only in the crown), Mo (Magnolia obovata only in the crown), Fc’ (periphery of F. crenata) and Gap. Seasonal changes in light availability above and below the Sasa layer was examined by using hemispherical photographs and quantum sensors.Results: Subcanopy individuals of F. crenata began unfolding their leaves approximately one month earlier than canopy trees of Q. mongolica var. grosseserrata and M. obovata, but a few days later than those of adult F. crenata. Accumulated photosynthetic photon flux density above the Sasa layer was greatest in Qm and Mo, and lowest in Fc. Importance of Sasa was highest in Gap. Maximum height and the number of subcanopy individuals of F. crenata were greatest in Qm, followed by Mo, and lowest in Fc.Conclusions: Differences in canopy layer composition probably influence the regeneration of F. crenata both directly through their foliage phenologies, and also indirectly by determining the importance of Sasa.

Influence of canopy tree phenology on understorey populations of Fagus crenata

Question: Is light available for subcanopy individuals of Fagus crenata spatiotemporally heterogeneous across patches with closed canopies of different foliage phenologies and gaps? Is local abundance of Sasa influenced by the composition of the canopy layer? If so, does the Sasa layer also affect the amount of light available to small F. crenata saplings? Is variation in F. crenata population structure consistent with the hypothesis that light is important? Location: Mt. Kurikoma, Japan 780 m a.s.l. Methods: Population structure of subcanopy individuals of Fagus crenata and importance of Sasa were examined in five patch types. The patch-types were Fc (F. crenata only in the crown), Qm (Quercus mongolica var. grosseserrata only in the crown), Mo (Magnolia obovata only in the crown), Fc’ (periphery of F. crenata) and Gap. Seasonal changes in light availability above and below the Sasa layer was examined by using hemispherical photographs and quantum sensors. Results: Subcanopy individuals of F. crenata began unfolding their leaves approximately one month earlier than canopy trees of Q. mongolica var. grosseserrata and M. obovata, but a few days later than those of adult F. crenata. Accumulated photosynthetic photon flux density above the Sasa layer was greatest in Qm and Mo, and lowest in Fc. Importance of Sasa was highest in Gap. Maximum height and the number of subcanopy individuals of F. crenata were greatest in Qm, followed by Mo, and lowest in Fc. Conclusions: Differences in canopy layer composition probably influence the regeneration of F. crenata both directly through their foliage phenologies, and also indirectly by determining the importance of Sasa.

Differences in properties of salt marsh sediment between hayed and reference sites.

than to seasonal changes in light availability. The increase of chlorophyll a and carotenoid concentrations in C. vagabunda and G. tikvahiae in response to high irradiance is paralleled by biomass data taken from the estuaries (E. Stieve, unpub. data). The biomass of G. tikvahiae was greater than that of C. vagabunda in low-nitrogen estuaries that also furnished high irradiance to algae. The physiological changes in photosynthetic pigments and nitrogen concentrations created by increased loads suggest increased growth of at least the green alga C. vagabunda. Increased nitrogen load may also increase phytoplankton standing crop, increasing light attenuation in the water column and therefore decreasing the growth of light-limited benthic algae. On balance, the growthstimulating effect of increased nutrients seems to more than compensate for the detrimental effect of light attenuation from the influence of phytoplankton shading at the time of sampling; and C. vagabunda, in particular, proliferates and causes macroalgal blooms in nitrogen enriched estuaries. This research was supported by the Woods Hole Marine Science Consortium. Special thanks to Jennifer Hauxwell for help with algae and Jennifer Bowen for her endless patience. Literature Cited

POPULATION DYNAMICS OF MICROALGAE IN THE UPPER LAND‐FAST SEA ICE AT A SNOW‐FREE LOCATION

The population dynamics of interior ice microalgae were investigated at a snow‐free site on annual land‐fast sea ice in McMurdo Sound, Antarctica, during the austral spring and summer of 1995–96. A dynamic successional sequence was observed with life history transformations playing an important role. During late November and early December (austral spring), cryo‐ and halotolerant dinoflagellates and chrysophytes bloomed in brine channels within the upper ice. At this time, competition and grazing pressure are low because of the inability of most marine species to grow under the extreme environmental conditions found in the upper ice during the austral spring. In November and December, dinoflagellates, chrysophytes, and prasinophytes contributed an average of 66%, 44%, and <1% of the phytoflagellate biomass, respectively. Both the dinoflagellates and the chrysophytes encysted in December, with cyst formation most intense just prior to surface melt and flushing of the ice. The cysts appear to be an adaptation for survival and dispersal in the plankton during ice decay and/or overwintering in the sea ice. In January (austral summer), when ice temperatures were similar to those in the water column, pennate diatoms replaced flagellates as the photosynthetic dominants in the upper sea ice. The upper land‐fast sea ice undergoes dramatic seasonal changes in light availability, temperature, brine salinity, and inorganic nutrient availability. Ephemeral blooms of cyst‐forming phytoflagellates exploit this habitat in the austral spring, when both inorganic nutrients and light are available but temperatures <−2° C and brine salinities elevated.

Organic Matter Budgets for Streams: A Synthesis

Organic Matter Budgets for Streams: A Synthesis