Photosynthetic acclimation of higher plants to growth in fluctuating light environments

This paper describes a study into the potential of plants to acclimate to light environments that fluctuate over time periods between 15 min and 3 h. Plants of Arabidopsis thaliana (L.) Heynh., Digitalis purpurea L. and Silene dioica (L.) Clairv. were grown at an irradiance 100 mol m^-2 s^-1. After 4–6 weeks, they were transferred to light regimes that fluctuated between 100 and either 475 or 810 mol m^-2 s^-1, in a regular cycle, for 7 days. Plants were shown, in most cases, to be able to undergo photosynthetic acclimation under such conditions, increasing maximum photosynthetic rate. The extent of acclimation varied between species. A more detailed study with S. dioica showed that this acclimation involved changes in both Rubisco protein and cytochrome f content, with only marginal changes in pigment content and composition. Acclimation to fluctuating light, at the protein level, did not fully reflect the acclimation to continuous high light - Rubisco protein increased more than would be expected from the mean irradiance, but less than expected from the high irradiance; cytochrome f increased when neither the mean nor the high irradiance would be expected to induce an increase.This revised version was published online in October 2005 with corrections to the Cover Date.     

Widespread correlations between dominance and homozygous effects of mutations: implications for theories of dominance

The dominance of deleterious mutations has important consequences for phenomena such as inbreeding depression, the evolution of diploidy, and levels of natural genetic variation. Kacser and Burns' metabolic theory provides a paradigmatic explanation for why most large-effect mutations are recessive. According to the metabolic theory, the recessivity of large-effect mutations is a consequence of a diminishing-returns relationship between flux through a metabolic pathway and enzymatic activity at any step in the pathway, which in turn is an inevitable consequence of long metabolic pathways. A major line of support for this theory was the demonstration of a negative correlation between homozygous effects and dominance of mutations in Drosophila, consistent with a central prediction of the metabolic theory. Using data on gene deletions in yeast, we show that a negative correlation between homozygous effects and dominance of mutations exists for all major categories of genes analyzed, not just those encoding enzymes. The relationship between dominance and homozygous effects is similar for duplicated and single-copy genes and for genes whose products are members of protein complexes and those that are not. A complete explanation of dominance therefore requires either a generalization of Kacser and Burns' theory to nonenzyme genes or a new theory.     

Measurement of in situ growth rates of phytoplankton under conditions of simulated turbulence

In situ measurement of the growth rates of planktonic populationscan be improved by using dialysis chambers (‘cage cultures’)to avoid shifts in the chemical environment during incubation.Vertical mixing and small-scale turbulence affect the growthof planktonic populations, there fore natural mixing conditionsshould be simulated as closely as possible during the incubation.A new device is described here which combines the advantagesof a dialysis chamber with a programmable vertical mixing regime.Realistic phytoplankton growth rates can thus be measured insitu under con ditions of vertical mixing and small-scale turbulence.The chamber made of transparent, UV-transmitting acrylic glasswas fitted at both ends with permeable polycarbonate membranes.It was moved vertically through the water column by a pocket-sizedlift and rotated simultaneously on its central axis. The methodwas applied to two typos of experiments on growth and lossesof phytoplankton in the River Severn, UK. The first one comparedchanges in biovolume of phytoplankton in a water parcel flowingdownstream (6% h^–1 decline) with those in a simultaneouslyincubated dialysis chamber moved between water surface and riverbottom (7% h^–1 increase). The difference equates to algallosses prevented in the chamber but suffered along the river(mainly sedimentation and grazing of benthic filter feeders).Loss rate of diatoms was three times higher than those of chlorophytes.In another experiment growth of phytoplankton from the mainstream and lateral dead zone was compared under different mixingconditions. Algae from the main stream grew faster than fromthe dead zone. Only cryptophytes preferred calm conditions,all the other algal groups grew faster in chambers moved throughthe water column than in stationary ones. Further possible applicationsin both standing and flowing waters are discussed.     

Unusual cleavage and gastrulation in a freshwater nematode: developmental and phylogenetic implications

Early embryogenesis in nematodes as seen in Caenorhabditis elegans and many other species of this phylum features several characteristic events. These include the visible presence of a germline from the very beginning generating different somatic lineages via asymmetric cleavages, the absence of a coeloblastula and a unique type of gastrulation with immigration of just two gut precursor cells. Here it is shown by using Nomarski optics that development of the freshwater nematode Tobrilus diversipapillatus differs from this pattern in two prominent aspects. (1) No asymmetric cleavages and no distinct cell lineages are generated; (2) in contrast to all other nematodes studied so far, a prominent coeloblastula is formed and gastrulation resembles the classical pattern found all over the animal kingdom. These developmental peculiarities are considered to be plesiomorphic and thus the order Triplonchida, to which Tobrilus belongs, may occupy a phylogenetic position at the base of the nematode phylum. The findings reported here allow us to reject a number of conceivable correlations between the type of gastrulation and other developmental parameters.Edited by D. TautzThis revised version was published online in December 2004 with corrections to typographical errors in the abstract.     

Correcting for allometry in studies of fluctuating asymmetry and quality within samples

Fluctuating asymmetry (FA) refers to deviations from perfect symmetry in a bilateral character and is believed to reflect an organism''s quality. However, allometric relations between asymmetry and trait size may confound FA–quality relations. Larger traits may have more ''opportunity'' to become asymmetric. Thus, researchers suggest that if allometric relations between FA and size are found, some correction for size is needed (typically FA scaled by size). I used a simulation model to examine potential consequences of allometry for detection of FA–quality relations within samples and examined efficacy of rules currently used to correct for size. Consequences of allometry can be severe, causing up to 77% type I errors or 98% power loss when true FA–quality relations exist (when FA–size relations are isometric), depending upon the strength and direction of size–quality relations. I illustrate why current rules to correct for size are inefficient and suggest alternative rules. Problems of allometry that require further attention are discussed.     

The effects of light and temperature on photosynthate partitioning in Antarctic freshwater phytoplankton

The effects of temperature and radiation flux on the partitioningof photosynthetically fixed carbon into four intracellulai metabolicpools was investigated for natural phytoplankton assemblagesfrom an Antarctic freshwater lake. At ambient temperature, proteinsynthesis was saturated at low photon flux densities (30–40µmol m^–2 s^–1) and above this flux fixed carbonwas increasingly stored as lipid and polysaccharide. Increasingtemperature raised both the saturated rate of protein synthesisand the photon flux at which saturation occurred. There wasa corresponding decline in the accumulation of reserve products,particularly at low radiation fluxes. The consequences of thispattern of uptake for the phytoplankton is discussed.     

Interactions between allelopathic properties and growth kynetics in four freshwater phytoplankton species studied by model simulations

We chose four species of freshwater phytoplankton: the chlorophyceans Ankistrodesmus falcatus, Chlamydomonas reinhardtii and Selenastrum capricornutum, and the cyanobacteria Oscillatoria sp. in order to study their competitive abilities for nitrate and their allelopathic properties. We parameterized models of nitrate uptake and growth with laboratory experiments. According to them, the species were ranked (from the best to the worst competitors): S. capricornutum, C. reinhardtii, A. falcatus and Oscillatoria sp. C. reinhardtii and Oscillatoria sp. were previously reported as allelopathic. In the present work, Oscillatoria sp. was allelopathic only against A. falcatus. However, none of our species was sensitive to C. reinhardtii. Additionally, we found an unknown allelopathic effect of A. falcatus against Oscillatoria sp. Our findings point out the high specificity of allelopathic interactions. With these data, we constructed a model of interspecific competition for nitrate, including allelopathic interactions. By performing model simulations, we studied how three factors influence the outcome of competition: relative abundance of competing species, resistance to allelopathy, and nitrate concentration. Our simulations showed that the initial ratio of species abundances will significantly determine the outcome of competition. If the worst competitor was the allelopathic species, the more it needs to outnumber the competing species, unless it is very sensitive to allelopathy (not defended). Nitrate has an important influence, showing a non-intuitive outcome of competition experiments at low nitrate concentrations, where the worst competitor (allelopathic species) wins competition in the majority of cases, whereas at intermediate concentrations, the better competitor dominates except for unfavorable ratios of abundances. With the increased amounts of nitrate, conditions again favor the worst competitor (the stronger allelopathic species). Despite the potential for two species coexistence showed by previous theoretical analysis of systems was similar to ours, our simulations did not detect this outcome. We hypothesized that this is due to the strong allelopathic effect of Oscillatoria sp.     

The allometry of fluctuating asymmetry in southern African plants: flowers and leaves

Several studies of fluctuating asymmetry (FA) in animals show that secondary sexual characters used in signalling have a negative relationship between size and asymmetry. Larger sexual traits are presumably more costly to produce, which should lead to greater developmental stress and corresponding increases in asymmetry. In the absence of among individual variation in the ability to handle these costs, the relationship between size and asymmetry should thus be positive. A negative relationship therefore suggests that expression of these traits is condition-dependent. In plants, flowers act as signals for pollinators and may show similar trends to animal signals. Leaves which are uninvolved in signalling should not. Moller & Eriksson (1994) found that 89% of species ( n = 16 of 18) with insect-pollinated flowers showed a negative relationship between petal size and asymmetry, while 79% of species ( n = 15 of 19) showed a positive relationship between leaf size and asymmetry. I carried out a similar study of 18 plant species. The average relationship between petal size and asymmetry did not differ significantly from zero in those species showing measurable FA in flowers ( n = 12). The relationship was significantly negative in one species, and significandy positive in another. On average, leaves in species with FA did not show a significant positive relationship between size and asymmetry ( n = 7). There was no significant difference in the slopes of the relationship between size and asymmetry for leaves and flowers. Levels of floral asymmetry for species with FA were significandy repeatable on individual plants in 33% ( n = 4 of 12) of species, but leaf asymmetry was not significantly repeatable in any species. It is argued that condition-dependence of traits need not result in a negative relationship between size and asymmetry.     

Plasticity of size and growth in fluctuating thermal environments: comparing reaction norms and performance curves

Ectothermic animals exhibit two distinct kinds of plasticityin response to temperature: Thermal performance curves (TPCs),in which an individual's performance (e.g., growth rate) variesin response to current temperature; and developmental reactionnorms (DRNs), in which the trait value (e.g., adult body sizeor development time) of a genotype varies in response to developmentaltemperatures experienced over some time period during development.Here we explore patterns of genetic variation and selectionon TPCs and DRNs for insects in fluctuating thermal environments.First, we describe two statistical methods for partitioningtotal genetic variation into variation for overall size or performanceand variation in plasticity, and apply these methods to availabledatasets on DRNs and TPCs for insect growth and size. Our resultsindicate that for the datasets we considered, genetic variationin plasticity represents a larger proportion of the total geneticvariation in TPCs compared to DRNs, for the available datasets.Simulations suggest that estimates of the genetic variationin plasticity are strongly affected by the number and rangeof temperatures considered, and by the degree of nonlinearityin the TPC or DRN. Second, we review a recent analysis of fieldselection studies which indicates that directional selectionfavoring increased overall size is common in many systems—thatbigger is frequently fitter. Third, we use a recent theoreticalmodel to examine how selection on thermal performance curvesrelates to environmental temperatures during selection. Themodel predicts that if selection acts primarily on adult sizeor development time, then selection on thermal performance curvesfor larval growth or development rates is directly related tothe frequency distribution of temperatures experienced duringlarval development. Using data on caterpillar temperatures inthe field, we show that the strength of directional selectionon growth rate is predicted to be greater at the modal (mostfrequent) temperatures, not at the mean temperature or at temperaturesat which growth rate is maximized. Our results illustrate someof the differences in genetic architecture and patterns of selectionbetween thermal performance curves and developmental reactionnorms.     

Dependence of phytoplankton assimilation quotients on light and nitrogen source: implications for oceanic primary productivity

Photosynthetic production of oxygen by phytoplankton assemblagedominated by Peridinium in Lake Kinneret, Israel, generallyexceeds the molar equivalent rate of carbon assimilation. Carbonassimilation occurs only if oxygenic photosynthesis exceedsa light-dependent threshold. Assimilation quotients (mol C molO₂^–1) are a variable function of irradiance, and typicallyonly about one-half of the photoreductant produced during oxygenicphotosynthesis is used for reduction of carbon dioxide. Mostof the residual oxygenic photoreductant probably is used forlight-dependent reduction of nitrate, which competes with carbondioxide for oxygenic photoreductant. Nitrate is an importantsource of nitrogen for this algal assemblage, and light-dependentnitrate reduction probably is much larger than carbon dioxidereduction at lowest irradiances in the euphotic zone. Oxygenproduction also may be much larger than carbon assimilationat low light levels in other environments where oxidized formsof nitrogen are important nitrogenous nutrients for phytoplankton,as in the lower euphotic zone of the sea, where low rates ofcarbon assimilation by phytoplankton have been thought to beinconsistent with the amount of oxygen that accumulates duringsummer.     

Critical flow velocities for the growth and dominance of Anabaena circinalis in some turbid freshwater rivers

SUMMARY 1. From measurements at several weir pool sites along the turbid and freshwater Barwon‐Darling River, Australia, the development of persistent stratification (for periods of >5 days) was related to river discharge. For the sites examined, the required discharge to allow the development of persistent stratification was between 100 and 450 ML day^?1 during the hotter months. High discharge during the hotter months did not allow the formation of persistent stratification, although diel stratification did occur. Low discharge through the cooler months resulted in diel stratification, although persistent stratification lasting for a few days could occur at times. 2. The growth and dominance of Anabaena circinalis at these sites was closely related to the establishment and maintenance of persistent and strong thermal stratification. Growth only occurred during extended periods (>5 days) of persistent stratification. These conditions not only restrict the displacement of A. circinalis downstream, they also allowed the alga to accumulate in surface waters. 3. The discharge levels required to suppress the formation of persistent stratification at the study sites were variable because of large differences in channel cross‐sectional area. To compensate for this variation, the discharges were converted to flow velocities. A critical velocity of 0.05 ms^?1 was sufficient for the suppression of persistent thermal stratification and concurrent A. circinalis growth for all sites. The turbulent velocity (u*) under weak wind mixing at the study locations varied between 2.66 × 10^?3 and 2.91 × 10^?3 ms^?1 at the critical flow velocities. These values may have potential to be applied to other rivers in similar climatic zones to suppress nuisance cyanobacterial growth.     

Response of lake phytoplankton communities to in situ manipulations of light intensity and colour

Phytoplankton preferences for light intensity and colour weredetermined in field experiments using coloured plexiglass cubessuspended at different depths in Heney Lake, Québec.Diatoms and green algae favoured intensities greater than 1%I_o (surface irradiance) contrary to dinoflagellates and otherflagellates that preferred lower intensity. Red radiation usuallyincreased the relative proportion of blue-greens, diatoms andgreen algae, whereas it reduced that of dinoflageilates. Wepropose that differential utilization of the light gradientallows certain phytoplankton taxa to partition the water column,thereby reducing potential competition. This is supported bythe general agreement between our findings and the known depthdistribution of algae in lakes.     

In situ growth rates of marine phytoplankton: approaches to measurement, community and species growth rates

Taxonomic structure and biomass weighting are important determinantsof measurable marine phytoplankton community growth potential.Maximal diel-averaged growth rates of communities appear tofall between 3 and 3.6 doublings day^–1. Mean net growthrates are considerably lower. Ranges of community growth ratesmeasured in tropical, sub-tropical and (summer) temperate ecosystemsare similar. There appears to be a broad dichotomy between thegrowth potential of diatom species as compared to non-diatoms.Doubling rates of small diatoms frequently exceed communitybiomass doubling rates by wide margins. Diel-averaged growthrates of large diatoms, microflagellates and non-motile ultraplanktonpopulations are lower and similar in magnitude to communitygrowth estimates. Maximum growth rates of species measured insitu are in good agreement with maximum growth rates measuredin laboratory cultures. High specific productivity of sub-dominantor rare diatom species or assemblages will be diluted in thelower specific growth rates of microflagellate and non-motileultraplankton assemblages. Specific rates of grazing upon speciesand functional groups remain to be quantified, but stabilityof community size and taxonomic structure implies close linkagebetween growth and mortality rates at the species level overtime intervals of several generations.     

Phytochrome control of the growth and development of Rumex obtusifolius under simulated canopy light environments*

Abstract Studies on the growth and development of Rumex obtusifolius seedlings under simulated shade conditions show that both light quantity and quality contribute to the observed responses. In the shade situation the plants have lower dry matter, lower leaf area and lower net assimilation rates. Petiole elongation occurs under shade conditions only after transfer of the plants from non-shade environments. The effects of light quality are related to phytochrome photoequilibria set up by the increased relative photon flux in the far-red found under vegetation canopies.     

Genetic recombination and adaptation to fluctuating environments: selection for geotaxis in Drosophila melanogaster

Heritable variation in fitness is the fuel of adaptive evolution, and sex can generate new adaptive combinations of alleles. If the generation of beneficial combinations drives the evolution of recombination, then the level of recombination should result in changes in the response to selection. Three types of lines of Drosophila melanogaster varying in their level of genetic recombination were selected over 38 generations for geotaxis. The within-chromosome recombination level of these lines was controlled for 60% of the genome: chromosome X and chromosome II. The full recombination lines had normal, unmanipulated levels of recombination on these two chromosomes. Conversely, nonrecombination lines had recombination effectively eliminated within the X and second chromosomes. Finally, partial recombination lines had the effective rate of within-chromosome recombination lowered to 10% of natural levels for these two chromosomes. The rate of response to selection was measured for continuous negative geotaxis and for a fluctuating environment (alternating selection for negative and positive geotaxis). All selected Drosophila lines responded to selection and approximately 36% of the response to selection was because of the X and second chromosomes. However, recombination did not accelerate adaptation during either directional or fluctuating selection for geotaxis.     

Optical and hydrographic consequences of freshwater run-off during spring phytoplankton growth in a Scottish fjord

A combination of in situ measurements and radiative transfermodelling were used to study optical conditions in the innerbasin of Loch Etive, a Scottish fjord, in March and April 2000.The basin was strongly stratified with three layers separatedby marked pycnoclines. The surface layer averaged 5 m in depthand was heavily stained with coloured dissolved organic matter(CDOM) which reduced the euphotic depth to between 7 and 10m. Approximately 20% of the photosynthetically available radiation(PAR) in the water column was absorbed by phytoplankton, 44%by CDOM and 36% by sea water. Detectable concentrations of themajor inorganic nutrients (nitrate, phosphate and silicate)occurred at all depths, but significant phytoplankton populations(averaging 6 mg chlorophyll a m^–3) were found only inthe reduced-salinity surface layer. The freshwater input thereforeacted both as a source of buoyancy which promoted phytoplanktongrowth near the surface and as an attenuator of visible lightwhich inhibited growth deeper in the water column.     

Developmental causes of allometry: new models and implications for phenotypic plasticity and evolution

Shapes change during development because tissues, organs, and various anatomical features differ in onset, rate, and duration of growth. Allometry is the study of the consequences of differences in the growth of body parts on morphology, although the field of allometry has been surprisingly little concerned with understanding the causes of differential growth. The power-law equation y?=?ax(b), commonly used to describe allometries, is fundamentally an empirical equation whose biological foundation has been little studied. Huxley showed that the power-law equation can be derived if one assumes that body parts grow with exponential kinetics, for exactly the same amount of time. In life, however, the growth of body parts is almost always sigmoidal, and few, if any, grow for exactly the same amount of time during ontogeny. Here, we explore the shapes of allometries that result from real growth patterns and analyze them with new allometric equations derived from sigmoidal growth kinetics. We use an extensive ontogenetic dataset of the growth of internal organs in the rat from birth to adulthood, and show that they grow with Gompertz sigmoid kinetics. Gompertz growth parameters of body and internal organs accurately predict the shapes of their allometries, and that nonlinear regression on allometric data can accurately estimate the underlying kinetics of growth. We also use these data to discuss the developmental relationship between static and ontogenetic allometries. We show that small changes in growth kinetics can produce large and apparently qualitatively different allometries. Large evolutionary changes in allometry can be produced by small and simple changes in growth kinetics, and we show how understanding the development of traits can greatly simplify the interpretation of how they evolved.     

Photosynthetic regulation under fluctuating light in field-grown Cerasus cerasoides: A comparison of young and mature leaves

Photosystem I (PSI) is a potential target of photoinhibition under fluctuating light. However, photosynthetic regulation under fluctuating light in field-grown plants is little known. Furthermore, it is unclear how young leaves protect PSI against fluctuating light under natural field conditions. In the present study, we examined chlorophyll fluorescence, P700 redox state and the electrochromic shift signal in the young and mature leaves of field-grown Cerasus cerasoides (Rosaceae). Within the first seconds after any increase in light intensity, young leaves showed higher proton gradient (ΔpH) across the thylakoid membranes than the mature leaves, preventing over-reduction of PSI in the young leaves. As a result, PSI was more tolerant to fluctuating light in the young leaves than in the mature leaves. Interestingly, after transition from low to high light, the activity of cyclic electron flow (CEF) in young leaves increased first to a high level and then decreased to a stable value, while this rapid stimulation of CEF was not observed in the mature leaves. Furthermore, the over-reduction of PSI significantly stimulated CEF in the young leaves but not in the mature leaves. Taken together, within the first seconds after any increase in illumination, the stimulation of CEF favors the rapid lumen acidification and optimizes the PSI redox state in the young leaves, protecting PSI against photoinhibition under fluctuating light in field-grown plants.     

An in situ pulse light fluorometer for chlorophyll determination as a monitor for vertical and horizontal phytoplankton distribution in lakes

Methodological problems with in vivo, fluorescence (IVF) measurementusing an in situ pulse light fluorometer were investigated inorder to validate this method for monitoring the vertical andhorizontal chlorophyll a (chl. a) distribution in lakes. Thecorrelation between chl. a and IVF was poor in the upper epilimnion(0–5 m) of a mesotrophic lake. The IVF of algal culturesand natural phytoplankton is very sensitive to changes in thelight environment. The response of the IVF to rapid light fluctuationsdepends on the amplitude of the light intensity and the lightconditions to which the algae were exposed before the onsetof light fluctuations. The variability of the ratio IVF:chl.a concentration makes a permanent calibration of the IVF againstchl. a necessary. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP), Second International Workshop heldat the National Oceanographic Institute, Haifa, Israel in April–May1984.     

Accuracy of circadian entrainment under fluctuating light conditions: contributions of phase and period responses.

The accuracy with which a circadian pacemaker can entrain to an environmental 24-h zeitgeber signal depends on (a) characteristics of the entraining signal and (b) response characteristics and intrinsic stability of the pacemaker itself. Position of the sun, weather conditions, shades, and behavioral variations (eye closure, burrowing) all modulate the light signal reaching the pacemaker. A simple model of a circadian pacemaker allows researchers to explore the impact of these factors on pacemaker accuracy. Accuracy is operationally defined as the reciprocal value of the day-to-day standard deviation of the clock times at which a reference phase (0) is reached. For the purpose of this exploration, the authors used a model pacemaker characterized solely by its momentary phase and momentary velocity. The average velocity determines the time needed to complete one pacemaker cycle and, therefore, is inversely proportional to pacemaker period. The model pacemaker responds to light by shifting phase and/or changing its velocity. The authors assumed further that phase and velocity show small random fluctuations and that the velocity is subject to aftereffects. Aftereffects were incorporated mathematically in a term allowing period to contract exponentially to a stable steady-state value, with a time constant of 69 d in the absence of light. The simulations demonstrate that a pacemaker reaches highest accuracy when it responds to light by simultaneous phase shifts and changes of its velocity. Phase delays need to coincide with slowing down and advances with speeding up; otherwise, no synchronization to the zeitgeber occurs. At maximal accuracy, the changes in velocity are such that the average period of the pacemaker under entrained conditions equals 24 h. The results suggest that during entrainment, the pacemaker adjusts its period to 24 h, after which daily phase shifts to compensate for differences between the periods of the zeitgeber and the clock are no longer necessary. On average, phase shifts compensate for maladjustments of phase and velocity changes compensate for maladjustments of period.