128 Capturing the Light Fantastic: Oceanographic and Tidal Influences on Intertidal Algae

Seaweeds experience many challenges to their persistence in intertidal zone habitats. Their growth rates must exceed losses associated with a range of ecological and physiological factors including desiccation, herbivory and wave forces. Growth rates depend on an alga's ability to capture and process light to build carbon-based molecules. We examined local (tidal height) and large (oceanographic) scale influences on algal photosynthetic efficiency and light climate, respectively. At the local scale, we combined periodic measurements of physiological state using PAM fluorometry with traditional demographic monitoring of a Postelsia palmaeformis , population over a tidal height gradient. Parameter estimates derived from rapid fluorescence-irradiance curves were correlated with longer-term ecological performance measures including growth rate, morphology, survivorship and reproductive output. At larger scales, we made continuous in situ measurements of chlorophyll fluorescence and light attenuation in the intertidal zone at six sites during 2001 and 2002. Light attenuation to the benthos was sharply reduced at sites when chl-a fluorescence was high. Long-term, large-scale monitoring of intertidal zone chl-a and macroalgal abundances documents that striking differences among sites are persistent and associated with oceanographic factors. The light saturation parameter and maximum photosynthetic rate calculated for several common intertidal macrophytes, along with published values of the irradiance needed to saturate their growth rates, suggest that underwater light levels may limit intertidal algal growth where phytoplankton blooms are common and persistent. We conclude that physiological stress associated with tidal and oceanographic factors contribute to macroalgal distributions.     

A simulation model of tree architecture development based on growth response to local light environment

A new model of three-dimensional tree architecture development was made, in which the growth of branches depends on their local light environment. The unit of the tree architecture is a linear stem called the branch unit (BU). Current-year BU's have leaves at their distal end. The local light environment is calculated considering mutual shading among leaved BU's. During the growth of a model tree, the number of leaved BU's increases and mutual shading becomes severe. The shadling leads to production of fewer new BU's and the death of some BU's, both of which restrain the overcrowding of BU's. The shape of the crowns of trees grown in a model forest stand varies with their position in the stand in a similar way as observed in real forests. This also results from the growth response of BU's to their local light environment. A model tree in which the photoassimillates were shared equally among the BU's was much disadvantaged in competition with the original model trees.     

A light distribution model for an internally radiating photobioreactor

Analysis of light energy distribution in culture is important for maximizing the growth efficiency of photosynthetic cells and the productivity of a photobioreactor. To characterize the irradiance conditions in a photobioreactor, we developed a light distribution model for a single-radiator system and then extended the model to multiple radiators using the concept of parallel translation. Mathematical expressions for the local light intensity and the average light intensity were derived for a cylindrical photobioreactor with multiple internal radiators. The proposed model was used to predict the irradiance levels inside an internally radiating photobioreactor using Synechococcus sp. PCC 6301 as a model photosynthetic microorganism. The effects of cell density and radiator number were interpreted through photographic and model simulation studies. The predicted light intensity values were found to be very close to those obtained experimentally, which suggests that the proposed model is capable of accurately interpreting the local light energy profiles inside the photobioreactor system. Due to the simplicity and flexibility of the proposed model, it was also possible to predict the light conditions in other complex photobioreactors, including optical-fiber and pond-type photobioreactors.     

木荷种源间光合作用参数分析

基于直角双曲线修正模型估算3个不同纬度的木荷种源(开平、太平和永丰种源)光补偿点、饱和点、最大净光合速率等参数,以便为评价不同木荷种源对环境的适应能力和优良种源选择等提供科学依据。结果表明:在3个不同纬度种源中,开平种源具有最高的净光合速率、最大净光合速率和较高的表观量子效率,且其生长速率最快;太平种源光饱和点最高,但其净光合速率、表观量子效率和最大净光合速率最低,其生长速率也最低;永丰种源具有较高的净光合速率、最大净光合速率和表观量子效率,其生长速率略高于太平种源。3个不同纬度木荷种源中,南部开平种源对当地环境具有较强的适应能力和生长潜力,具有较高的推广应用价值。     

Comparative growth ecology of Ascophyllum nodosum from Norway (lat. 63°N) and Wales (LAT. 53°N)

Two populations of Ascophyllum nodosum (L.) Le Jol., one from Menai Bridge, Wales, other from Trondheim, Norway, were grown in Trondheim under identical conditions. The two populations showed an equal growth response to light (saturation at 100 Wm⁻² at a day-length > 18 h). Temperature optima were 17–18°C and 16°C for the Menai Bridge and Trondheim populations, respectively.In Menai Bridge, local apical growth during April–August was about twice that in Trondheim. When the two populations were exposed to identical conditions (natural light, variable irradiances, different daylengths, different temperatures), the Menai Bridge population had about a 40% better growth rate than the Trondheim population, irrespective of treatment. It is concluded that the differences in local growth rates are partly due to environmental conditions and partly to different genetic growth capacities.     

FTIR spectra of algal species can be used as physiological fingerprints to assess their actual growth potential

Fourier transform infrared (FTIR) spectra were measured from cells of Microcystis aeruginosa and Protoceratium reticulatum, whose growth rates were manipulated by the availability of nutrients or light. As expected, the macromolecular composition changed in response to the treatments. These changes were species‐specific and depended on the type of perturbation applied to the growth regime. Microcystis aeruginosa showed an increase in the carbohydrate‐to‐protein ratio with decreased growth rates, under nutrient limitation, whereas light limitation induced a decrease of the carbohydrate‐to‐protein ratio with decreasing proliferation rates. The macromolecular pools of P. reticulatum showed a higher degree of compositional homeostasis. Only when the lowest light irradiance and nutrient availability were supplied, an increase of the carbohydrate‐to‐protein FTIR absorbance ratio was observed. A species‐specific partial least squares (PLS) model was developed using the whole FTIR spectra. This model afforded a very high correlation between the predicted and the measured growth rates, regardless of the growth conditions. On the contrary, the prediction based on absorption band ratios generally used in FTIR studies would strongly depend on growth conditions. This new computational method could constitute a substantial improvement in the early warning systems of algal blooms and, in general, for the study of algal growth, e.g. in biotechnology. Furthermore, these results confirm the suitability of FTIR spectroscopy as a tool to map complex biological processes like growth under different environmental conditions.     

Growth Rates of Marine Phytoplankton: Correlation with Light Absorption by Cell Chlorophyll a

To better predict plant production in the sea, it would be desirable to be able to calculate, from easily obtainable measurements at one sampling, the growth rate of the prevailing stock of phytoplankton. To this end growth rates, pigment composition, cell volume and cell surface area data were collected for several species of marine phytoplankton in logarithmic growth at 20–21°C and 0.07 cal/cm². min light intensity. Similar data for one species, Dunaliella tertiolecta, are given for several combinations of light intensity and temperature, and for another species, Ditylum brightwellii, grown in nitrogen deficiency. The problem of estimating growth rates of phytoplankton was divided into three parts: 1) variation of growth rate among diverse species and its relationship to light absorption by cell chlorophyll a: 2) variation in growth rate with light intensity; 3) variation in growth rate with temperature. An equation has been formulated for calculating growth rate which provides a more precise fit of the data than do equations for growth rate based upon cell surface/volume ratios or cell volume. The formulation is based upon light absorption by chlorophyll a. It allows for variations in the efficiency of utilization of light absorbed by chlorophyll a and the changes in chlorophyll a content resulting from light intensity and temperature differences. We do not attempt to predict variations in growth rate with photoperiod or spectral distribution, nor do we allow for light effects upon growth rate not mediated by photosynthesis, so the model is, at best, a rough approximation of reality.     

Diffusion-limited predator-prey dynamics in Euclidean environments: an allometric individual-based model

We claim that diffusion-limited rates of reaction can be an explanation for the altered population dynamics predicted by models incorporating local interactions and limited individual mobility. We show that the predictions of a spatially explicit, individual-based model result from reduced rates of predation and reproduction caused by limited individual mobility and patchiness. When these reduced rates are used in a mean-field model, there is better agreement with the predictions of the simulation model incorporating local interactions. We also explain previous findings regarding the effects of dimensionality on population dynamics in light of diffusion-limited reactions and Pólya random walks. In particular, we demonstrate that 3D systems are better "stirred" than 2D systems and consequently have a reduced tendency for diffusion-limited interaction rates.     

Concentrative nitrogen allocation to sun-lit branches and the effects on whole-plant growth under heterogeneous light environments

We investigated the nitrogen and carbohydrate allocation patterns of trees under heterogeneous light environments using saplings of the devil maple tree (Acer diabolicum) with Y-shaped branches. Different branch groups were created: all branches of a sapling exposed to full light (L-branches), all branches exposed to full shade (S-branches), and half of the branches of a sapling exposed to light (HL-branches) and the other half exposed to shade (HS-branches). Throughout the growth period, nitrogen was preferentially allocated to HL-branches, whereas nitrogen allocation to HS-branches was suppressed compared to L- and S-branches. HL-branches with the highest leaf nitrogen content (N_area) also had the highest rates of growth, and HS-branches with the lowest N_area had the lowest observed growth rates. In addition, net nitrogen assimilation, estimated using a photosynthesis model, was strongly correlated with branch growth and whole-plant growth. In contrast, patterns of photosynthate allocation to branches and roots were not affected by the light conditions of the other branch. These observations suggest that tree canopies develop as a result of resource allocation patterns, where the growth of sun-lit branches is favoured over shaded branches, which leads to enhanced whole-plant growth in heterogeneous light environments. Our results indicate that whole-plant growth is enhanced by the resource allocation patterns created for saplings in heterogeneous light environments.     

Light‐based Regeneration Niches: Evidence from 21 Dipterocarp Species using Size‐specific RGRs

A continuing challenge in tropical ecology is to explain the coexistence of large numbers of rain forest tree species. One possible coexistence mechanism is partitioning of the highly variable and dynamic forest light environment, in which species that grow better in one light treatment grow worse in another. To test whether species respond differently to the light environment, we estimated growth rates of 21 Dipterocarpaceae species from Malaysian Borneo grown in shade houses for 2 yr in three light treatments (0.3%, 3%, and 18% full sunlight). We made regular measurements of height, diameter, and aboveground biomass, enabling us to calculate growth rates for each response. We estimated size‐specific growth rates using nonlinear mixed‐effects models, as average relative growth rate was strongly size dependent. For all species, the greatest diameter growth rate was achieved in 18 percent full sunlight, whereas for five of the twenty‐one species, the greatest height growth rate was achieved in three percent full sunlight. We investigated correlations among growth rates in different light treatments, but no negative correlations were found, indicating that species growing well in one light treatment did not grow poorly in the others. There were substantial crossovers, however, in species ranks among the three light treatments, indicating that there was no single growth rate hierarchy common to all light treatments. The lack of a single consistent growth hierarchy across light treatments indicates that heterogeneity in the forest light environment could contribute to the maintenance of the diversity of Dipterocarpaceae found in lowland Bornean rain forests via light‐based regeneration niches.     

The snail <Emphasis Type="Italic">Potamopyrgus antipodarum</Emphasis> grows faster and is more active in the shade,independent of food quality

Ecological stoichiometry has advanced food web ecology by emphasising the importance of food quality over food quantity for herbivores. Here, we focus on the effects of abiotic factors such as nutrients and light (known to influence food quality) on grazer growth rates. As model organism we used the mudsnail Potamopyrgus antipodarum that is native to New Zealand but invasive elsewhere. In a stream channel experiment in New Zealand, we manipulated light (two levels) and nutrients (four levels) to achieve a range of primary producer carbon: nutrient ratios and added mudsnails (3 densities + ungrazed control) to 128 periphyton covered stream channels in a 2 × 4 × 4 full factorial design. We measured snail growth rate and activity, food quality and nutritional imbalance, to test the predictions that (1) less light and more nutrients increase periphyton food quality and thus snail growth rates, and (2) less crowding leads to higher food availability and thus higher snail growth rates. We found that snail growth rates were higher under low light than under high light intensities and this difference increased with increasing nutrient addition. These changes in growth rate were not mediated by food quality in terms of periphyton nutrient ratios. Furthermore, experimental treatments strongly affected snail behaviour. Snails grazed more actively in the low light treatments, and thus it is more likely that snail growth rates were directly affected by light levels, maybe as a result of innate predator avoidance behaviour or as a reaction to high UV intensities. We conclude that in our stream channels snail growth rate was limited by factors other than food quality and quantity such as UV exposure, algal defences or the relatively low ambient water temperature.     

An ESS for the Height of a Plant Population, or an Optimal Height for an Individual?—Rethinking Game-Theoretic Models for Plant Height

Plants only interact with neighbors over restricted distances, so local conditions are of great significance for plants. It is therefore important to consider spatial structure and neighborhood effects if we are to understand plants' strategies. We constructed a spatially-explicit, game theory model to explore optimal height growth at the individual-level. In the model, there is no ESS for height growth at the population level, because there is an “instantaneous” optimal height growth strategy for the individual plant that changes depending on the local light environment. The optimal strategy is plasticity in response to local conditions. Game-theoretic models for plant phenotypic traits should move from “mean-field approximations” towards explicit modeling of local interactions.     

Shedding light on plant competition: modelling the influence of plant morphology on light capture (and vice versa)

A plant's morphology is both strongly influenced by local light availability and, simultaneously, strongly influences this local light availability. This reciprocal relationship is complex, but lies at the heart of understanding plant growth and competition. Here, we develop a sub-individual-based simulation model, cast at the level of interacting plant components. The model explicitly simulates growth, development and competition for light at the level of leaves, branches, etc., located in 3D space. In this way, we are able to explore the manner in which the low-level processes governing plant growth and development give rise to individual-, cohort-, and community-level phenomena. In particular, we show that individual-level trade-offs between growing up and growing out arise naturally in the model, and robustly give rise to cohort-level phenomena such as self-thinning, and community processes such as the effect of ecological disturbance on the maintenance of biodiversity. We conclude with a note on our methodology and how to interpret the results of simulation models such as this one.     

THE EFFECTS OF LIGHT AND TEMPERATURE ON GROWTH RATES IN BOREAL-SUBARCTIC CRUSTOSE CORALLINES1

A number of boreal-subarctic crustose corallines were kept in natural seawater tanks at temperatures ranging from 0 to 19 C and, using fluorescent lamps at light intensities, ranging from 7 to 750 lux with periods of 8 and 14 hr/day. The resultant growth rates as a function of temperature and light are presented and discussed in relation to the ecology of the plants. All of the Lithothamnieae studied had growth maxima at temperatures from 9 to 15 C. Growth in these species showed little light dependence below 4–6 C, but had a strong light dependence at higher temperatures. The one Lithophyllum species examined gave a flatter growth-temperature curve than the Lithothamnieae and showed little light dependence. The effects of temperature variation, salinity, and current on growth rates were also examined and are discussed. It was found to be especially important in studying growth rates of crustose corallines to allow time for growth stabilization following temperature change. In general, growth was found to exhibit a hysteresis effect, increased rates with the raising of temperatures 5–10 C and decreased rates with lowering temperatures.     

Effects of potential PAR on shoot extension in juveniles of the main tree species in a Japanese temperate forest

We studied the relationship between potential photosynthetically active radiation (PAR), estimated from hemispherical photographs, and shoot extension rates of juveniles of 12 seral tree species. We distinguished between direct light site factor (DIR), diffuse light site factor (DIF), and gap light index (GLI). We used a log-linear model to relate growth to DIR, DIF and GLI. Potential PAR explained shoot growth rates significantly in 33 cases out of 36 (12 species× 3 PAR). DIF explained the shoot extension rates better than DIR and GLI for 10 of the 12 species. The mean values of maximum shoot extension rates of the tolerant, the intermediate and the intolerant species were 52.4 cm year^–1, 64.3 cm year^–1 and 87.7 cm year^–1, respectively. The maximum shoot extension rates of the tolerant and the intolerant species were recorded in about 50% DIF and more than 70% DIF, respectively. The minimum light level for seedling establishment of the intermediate and intolerant species was more than 20% DIF. We explained distribution characteristics and shoot extension of the species in relation to light in terms of shade tolerance. Predictions based on light-growth curves and maximum growth curves were similar to field observations made elsewhere, suggesting that these models may be useful to predict extension growth of juvenile trees in mixed species stands.     

On the crystallization of proteins

We report on theoretical and experimental work aimed at a systematic approach to the crystallization of proteins. Successful crystallization depends on the competition between the growth rates for compact three-dimensional structures and long-chain structures leading to an amorphous precipitate. Quasi-elastic light scattering was used to monitor the size and shape distribution of small aggregates in a model system (lysozyme) during the pre-nucleation stage. With the aid of a simple model, the line-width of the scattered light was used to predict whether crystals or an amorphous precipitate would result. Once visible crystals appeared, the lysozyme concentration near the crystal surface was monitored and the kinetic parameters for growth obtained. A peculiar self-limiting phenomenon causes crystals to stop growing after a certain size has been reached. When these terminal size crystals were cleaved, growth occurred at the surface until the original size was approximately restored.     

PHOTO‐ACCLIMATION RESPONSE OF BENTHIC STREAM ALGAE ACROSS EXPERIMENTALLY MANIPULATED LIGHT GRADIENTS: A COMPARISON OF GROWTH RATES AND NET PRIMARY PRODUCTIVITY1

The objective of this study was to measure the photo‐acclimation response of stream algae inhabiting thin biofilms across a range of light treatments comparing both growth rates and net primary productivity (NPP). Algae were grown on clay tiles incubated in artificial stream channels where light levels were manipulated by layering a neutral‐density shade cloth over each channel. We measured NPP and algal growth rates in the early stages of community development and compared assemblages that were either acclimated or unacclimated to a given light treatment. Algal growth rates did acclimate to light treatment, with saturation occurring at light levels that were substantially lower for the acclimated communities. Growth efficiency calculated from algal biovolume increased by a factor of 3. However, algal NPP showed a weaker photo‐acclimation response, with only a 30% increase in photosynthetic efficiency. Our results indicate that diatom‐dominated periphyton in thin biofilms are probably not light limited in many shaded streams with respect to growth rates, but are light limited with respect to NPP.     

Feasibility and limits of an orthotopic human colon cancer model in nude mice

We sought to develop an accurate colorectal cancer model in nude mice with stable local growth, tumor cell dissemination, and reproducible metastatic capacity. To this end, we orthotopically transplanted histologically intact human colorectal cancer tissue from 10 human patients into nude mice. After successful local tumor growth, tumor tissues were retransplanted as many as 9 times in serial passage. All specimens were transplanted using microsurgical techniques. Histologic, immunohistochemical, and polymerase chain reaction techniques were used to determine tumor growth rates and kinetics, development of regional lymph node and distant hepatic metastases, and the induction of minimal residual disease (MRD). Stable local tumor growth rates with variable growth kinetics were detected in 73.4% of all mice. The lymph node and hepatic metastasis rates were low, at 18.4% and 4.9%, respectively. MRD, as reflected by CK20 positivity of the bone marrow in animals with lymph node and hepatic metastases, was present in 22.2%. The orthotopic colorectal cancer model described here is feasible for the induction of reproducible local tumor growth but is limited by variable growth kinetics and the low rate of lymph node and hepatic metastases. Cytokeratin-positive cells indicative of MRD could be detected in the bone marrow of approximately 25% of the nude mice with metastases. The observed induction of MRD after orthotopic implantation of intact human colon cancer in animals with lymph node and hepatic metastases might be improved if established colon cancer cell lines were used.     

Microalgae cultivation in air-lift reactors: modeling biomass yield and growth rate as a function of mixing frequency

The slow development of microalgal biotechnology stems from the failure in the design of large-scale photobioreactors where light energy is efficiently utilized. Due to the light gradient inside the reactor and depending on the mixing properties, algae are subjected to certain light/dark cycles where the light period is characterized by a light gradient. These light/dark cycles will determine productivity and biomass yield on light energy. Air-lift reactors can be used for microalgae cultivation and medium-frequency light/dark cycles will be found in these systems. Light/dark cycles are associated with two basic parameters: first, the light fraction, i.e., the ratio between the light period and the cycle time and second, the frequency of the light/dark cycle. In the present work, light/dark cycles found in air-lift reactors were simulated taking into account the light gradient during the light period. The effect of medium-frequency cycle time (10-100 s) and light fraction (0.1-1) on growth rate and biomass yield on light energy of the microalgae Dunaliella tertiolecta was studied. The biomass yield and growth rates were mainly affected by the light fraction, while cycle time had little influence. Response surface methodology was used and a statistical model describing the effect of light fraction and cycle time on growth rate and biomass yield on light energy was developed. The use of the model as a reactor design criterion is discussed.     

What shapes local density? The importance of migration rates and local growth for density‐patch size relationships in two Cionus weevils

1. The relative effect of migration and local growth on the spatio‐temporal density‐distribution of two co‐existing herbivorous weevils, Cionus scrophulariae L. and C. tuberculosus Scop., in 32 host plant Scrophularia nodosa L. patches of varying sizes was investigated. 2. Predictions of the temporal development of the slope in the density‐patch size relationships were derived from a basic population model with scale‐dependent migration rates. The model indicated that the slopes in the density‐patch size relationships during the early season should be reflected by the net scaling of immigration and emigration rates, whereas the slopes during the later season should increase as a result of local growth. 3. Emigration rates of the weevils were estimated in a field experiment, were the weevils coexisted in space and time. These results were then combined with a previous estimate of immigration rates in order to determine the net scaling of migration rates. 4. The emigration rate differed between species, caused by different movement rates in small patches, which could explain differences in the general slope of the density‐patch size relationships of the weevils in the natural figwort patches throughout the summer. The slopes in the relationships in the early season were largely predicted by the net scaling of migration rates. The slope also increased in the later season for C. tuberculosus, whereas the slope decreased for C. scrophulariae. 5. It was concluded that the understanding of both inter‐ and intra‐specific variations in density‐patch size relationships of insect herbivores can be improved using population models incorporating scale‐dependent migration and local growth.