Low-light Habitats Research Articles

Minimal transcriptional regulation of horizontally transferred photosynthesis genes in phototrophic bacterium Gemmatimonas phototrophica.

Horizontal gene transfer is one of the main mechanisms by which bacteria acquire new genes. However, it represents only the first step as the transferred genes have also to be functionally and regulatory integrated into the recipient's cellular machinery. Gemmatimonas phototrophica, a member of bacterial phylum Gemmatimonadota, acquired its photosynthesis genes via distant horizontal gene transfer from a purple bacterium. Thus, it represents a unique natural experiment, in which the entire package of photosynthesis genes was transplanted into a distant host. We show that G. phototrophica lacks the regulation of photosynthesis gene expressions in response to oxygen concentration and light intensity that are common in purple bacteria. This restricts its growth to low-light habitats with reduced oxygen. Understanding the regulation of horizontally transferred genes is important not only for microbial evolution but also for synthetic biology and the engineering of novel organisms, as these rely on the successful integration of foreign genes.

Splitting light pollution: Wavelength effects on the activity of two sandy beach species

Artificial Light at Night (ALAN) threatens to disrupt most natural habitats and species, including those in coastal settings, where a growing number of studies have identified ALAN impacts. A careful examination of the light properties behind those impacts is important to better understand and manage the effects of this stressor. This study focused on ALAN monochromatic wavelengths and examined which types of light spectra altered the natural activity of two prominent coastal species from the Pacific southeast: the talitroid amphipod Orchestoidea tuberculata and the oniscoid isopod Tylos spinulosus. We compared the natural daylight/night activity of these organisms with the one they exhibit when exposed to five different ALAN wavelengths: lights in the violet, blue, green, amber, and red spectra. Our working hypothesis was that ALAN alters these species’ activity at night, but the magnitude of such impact differs depending on light wavelengths. Measurements of activity over 24 h cycles for five consecutive days and in three separate experiments confirmed a natural circadian activity pattern in both species, with strong activity at night (∼90% of probability) and barely any activity during daylight. However, when exposed to ALAN, activity declined significantly in both species under all light wavelengths. Interestingly, amphipods exhibited moderate activity (∼40% of probability) when exposed to red lights at night, whereas isopods shifted some of their activity to daylight hours in two of the experiments when exposed to blue or amber lights, suggesting a possible alteration in this species circadian rhythm. Altogether, our results were consistent with our working hypothesis, and suggest that ALAN reduces night activity, and some wavelengths have differential effects on each species. Differences between amphipods and isopods are likely related to their distinct adaptations to natural low-light habitat conditions, and therefore distinct sensitivity to ALAN.

The native stem holoparasitic Cuscuta japonica suppresses the invasive plant Ambrosia trifida and related mechanisms in different light conditions in northeast China.

Increasing evidence from low-latitude ranges has demonstrated that native parasitic plants are promising biocontrol agents for some major invasive weeds. However, related mechanisms and the effect of environments on the control effect of the parasite are still unclear. In addition, few related studies have been conducted in high latitude (>40°), where the exotic plant richness is the highest in the globe, but natural enemies are relatively scarce. During field surveys, a Cuscuta species was found on the cosmopolitan invasive weed Ambrosia trifida L. in Shenyang, northeast China. Here, we first studied the impacts of the parasite on the invader at three sites with different light regimes and related mechanisms, then the haustorial connections between the parasite and the invader using anatomy and measurement of carbon (C) and nitrogen (N) stable isotope compositions (δ13C, δ15N), and finally identified the parasite using two molecular marks. The parasite was identified as C. japonica Choisy. This native holoparasitic vine posed serious C rather than N limitation to the invader, explaining its greatly inhibitory effects on the invader. Its negative effects were stronger on reproductive relative to vegetative growth, and at high relative to low light habitats, which indicated that the higher the vigor of the host is, the higher the impact of the parasite pose. The parasite could establish haustorial connections with phloem, xylem, and pith of the invader and thus obtain resources from both leaves and roots, which was confirmed by difference of δ13C and δ15N between the two species. The parasite had significantly higher leaf C concentrations and δ13C than its invasive host, being a strong C sink of the parasitic association. Our results indicate that C. japonica may be a promising biological control agent for the noxious invader in China.

Harnessing solar power: photoautotrophy supplements the diet of a low-light dwelling sponge

The ability of organisms to combine autotrophy and heterotrophy gives rise to one of the most successful nutritional strategies on Earth: mixotrophy. Sponges are integral members of shallow-water ecosystems and many host photosynthetic symbionts, but studies on mixotrophic sponges have focused primarily on species residing in high-light environments. Here, we quantify the contribution of photoautotrophy to the respiratory demand and total carbon diet of the sponge Chondrilla caribensis, which hosts symbiotic cyanobacteria and lives in low-light environments. Although the sponge is net heterotrophic at 20 m water depth, photosynthetically fixed carbon potentially provides up to 52% of the holobiont’s respiratory demand. When considering the total mixotrophic diet, photoautotrophy contributed an estimated 7% to total daily carbon uptake. Visualization of inorganic 13C- and 15N-incorporation using nanoscale secondary ion mass spectrometry (NanoSIMS) at the single-cell level confirmed that a portion of nutrients assimilated by the prokaryotic community was translocated to host cells. Photoautotrophy can thus provide an important supplemental source of carbon for sponges, even in low-light habitats. This trophic plasticity may represent a widespread strategy for net heterotrophic sponges hosting photosymbionts, enabling the host to buffer against periods of nutritional stress.

Episkeletobionts of large rugose corals from the Middle Devonian mesophotic palaeoenvironment recorded in the Pokrzywianka Beds (Holy Cross Mountains, Poland)

Organisms encrusting corals from a coral horizon encountered in a trench in the Middle Devonian (Givetian) Pokrzywianka Beds of the classic Grzegorzowice-Skały section in the Holy Cross Mountains, Poland, are described and analyzed in the context of their palaeoecological and palaeoenvironmental background. These episkeletobionts form rather a low-diversity community, dominated by microconchid tubeworms, crinoids, and tabulate corals. The last group, however, is especially diverse at the family level, represented by auloporids, alveolitids, coenitids and favositids. These episkeletobionts are considered to have developed in a low-light, lower mesophotic palaeoeonvironment, as evidenced by the presence of platy, alveolitid tabulate coral in the deposits studied. This microconchid-crinoid-tabulate-coral community differs from other Givetian communities from the Holy Cross Mountains (Laskowa and Miłoszów), which also are considered to have developed in low-light habitats. The differences in taxonomic composition of episkeletobionts between these three localities most probably resulted from specific local conditions, related to bathymetry (light levels, nutrient levels), the specific nature of the hosts/substrates occupied, and also differences in larval dispersal patterns. This, in turn, shows that various encrusting communities may have inhabited seemingly similar, marine habitats within a given time interval and neighbouring areas, which may have serious implications for large-scale comparisons of biodiversity within a given palaeoenvironment.

Exploring coral reef biodiversity via underwater soundscapes

Information on biodiversity is essential to evaluate the ecological status of coral reefs. Sounds produced by reef-associated organisms have been used as a biodiversity indicator. However, the interference from abiotic sounds and the lack of a comprehensive audio library have impeded effective evaluation. This study investigated the application of underwater soundscapes as a remote-sensing method to detect biological and anthropogenic activities. Using techniques including the visualization of long-duration recordings, source separation, and clustering, soundscapes were separated into sounds of anthropogenic and biological sources. Our results revealed the dynamics of biological sounds among coral reefs off Sesoko Island, Okinawa, Japan. Biological sounds were much more prominent in shallow-water reefs than in upper-mesophotic reefs, but their spectral features and compositions differed. The shallow-water reefs were dominated by broadband sounds of crustaceans and low-frequency transient fish calls, whereas the upper-mesophotic reefs were characterized by a diverse array of fish choruses and transient sounds. We also discovered that shipping noise heavily interfered with the soundscapes from the upper-mesophotic reefs and represented an invisible threat to life in the low-light habitat. The applied techniques of soundscape information retrieval revealed the distinct ecological status of coral reefs and the behavior change of sound-producing organisms in high temporal resolution. Implementation of soundscape monitoring can generate ecological information on habitat quality, reef biodiversity, human activities, and their interactions. Global collaboration on underwater soundscapes will establish a data-informed platform and help stakeholders assess the resilience of coral reefs to environmental and anthropogenic stressors.

Genomic Analysis of the Only Blind Cichlid Reveals Extensive Inactivation in Eye and Pigment Formation Genes.

Trait loss represents an intriguing evolutionary problem, particularly when it occurs across independent lineages. Fishes in light-poor environments often evolve “troglomorphic” traits, including reduction or loss of both pigment and eyes. Here, we investigate the genomic basis of trait loss in a blind and depigmented African cichlid, Lamprologus lethops, and explore evolutionary forces (selection and drift) that may have contributed to these losses. This species, the only known blind cichlid, is endemic to the lower Congo River. Available evidence suggests that it inhabits deep, low-light habitats. Using genome sequencing, we show that genes related to eye formation and pigmentation, as well as other traits associated with troglomorphism, accumulated inactivating mutations rapidly after speciation. A number of the genes affected in L. lethops are also implicated in troglomorphic phenotypes in Mexican cavefish (Astyanax mexicanus) and other species. Analysis of heterozygosity patterns across the genome indicates that L. lethops underwent a significant population bottleneck roughly 1 Ma, after which effective population sizes remained low. Branch-length tests on a subset of genes with inactivating mutations show little evidence of directional selection; however, low overall heterozygosity may reduce statistical power to detect such signals. Overall, genome-wide patterns suggest that accelerated genetic drift from a severe bottleneck, perhaps aided by directional selection for the loss of physiologically expensive traits, caused inactivating mutations to fix rapidly in this species.

Photoacclimation in freshwater diatoms: interspecific and inter-habitat comparisons

Diatoms are a type of photosynthetic phytoplankton that can acclimate to the light level of their environment. Diatoms from different habitats exhibit different photoacclimation characteristics; those from relatively low-light intensity habitats present relatively high photosynthetic activity, high light-harvesting pigment (Chla) content, and low xanthophyll content. Conversely, diatoms from high-light intensity environments have lower photosynthetic activity, less Chla content, and efficient xanthophyll cycling. Hence, diatoms can maintain high growth and photosynthetic activity within a wide range of light intensities. Whether the mechanisms of such adaptability are a consequence of biological photoacclimation to different light intensities or interspecific differences is unknown. Here, we show that differences in photoacclimation ability are more significant than species differences. We found similar species compositions in low- and high-light habitats of the Hanjiang River, China; however, there were remarkable differences in rapid light-response curve parameters. The photoacclimation of algae and trends in their photosynthetic activity can be estimated by rapid detection techniques. We isolated three diatom species common to both habitats and grew them under various light intensities, finding that they have an excellent ability to acclimate to local light conditions. Diatoms can use physiological strategies to handle light fluctuations, but only for short periods. Our results provide a theoretical basis for controlling algal blooms through light management. With knowledge of the photoacclimation characteristics of diatoms and according to the availability of local water conservancy facilities, it is possible to use light management to control diatom blooms more efficiently than with conventional techniques, thereby reducing water usage.

De novo genome assembly of Oryza granulata reveals rapid genome expansion and adaptive evolution

The wild relatives of rice have adapted to different ecological environments and constitute a useful reservoir of agronomic traits for genetic improvement. Here we present the ~777 Mb de novo assembled genome sequence of Oryza granulata. Recent bursts of long-terminal repeat retrotransposons, especially RIRE2, led to a rapid twofold increase in genome size after O. granulata speciation. Universal centromeric tandem repeats are absent within its centromeres, while gypsy-type LTRs constitute the main centromere-specific repetitive elements. A total of 40,116 protein-coding genes were predicted in O. granulata, which is close to that of Oryza sativa. Both the copy number and function of genes involved in photosynthesis and energy production have undergone positive selection during the evolution of O. granulata, which might have facilitated its adaptation to the low light habitats. Together, our findings reveal the rapid genome expansion, distinctive centromere organization, and adaptive evolution of O. granulata.

The Toxicogenome of Hyalella azteca: A Model for Sediment Ecotoxicology and Evolutionary Toxicology.

Hyalella azteca is a cryptic species complex of epibenthic amphipods of interest to ecotoxicology and evolutionary biology. It is the primary crustacean used in North America for sediment toxicity testing and an emerging model for molecular ecotoxicology. To provide molecular resources for sediment quality assessments and evolutionary studies, we sequenced, assembled, and annotated the genome of the H. azteca U.S. Lab Strain. The genome quality and completeness is comparable with other ecotoxicological model species. Through targeted investigation and use of gene expression data sets of H. azteca exposed to pesticides, metals, and other emerging contaminants, we annotated and characterized the major gene families involved in sequestration, detoxification, oxidative stress, and toxicant response. Our results revealed gene loss related to light sensing, but a large expansion in chemoreceptors, likely underlying sensory shifts necessary in their low light habitats. Gene family expansions were also noted for cytochrome P450 genes, cuticle proteins, ion transporters, and include recent gene duplications in the metal sequestration protein, metallothionein. Mapping of differentially expressed transcripts to the genome significantly increased the ability to functionally annotate toxicant responsive genes. The H. azteca genome will greatly facilitate development of genomic tools for environmental assessments and promote an understanding of how evolution shapes toxicological pathways with implications for environmental and human health.

Varying and unchanging whiteness on the wings of dusk-active and shade-inhabiting Carystoides escalantei butterflies

Whiteness, although frequently apparent on the wings, legs, antennae, or bodies of many species of moths and butterflies, along with other colors and shades, has often escaped our attention. Here, we investigate the nanostructure and microstructure of white spots on the wings of Carystoides escalantei, a dusk-active and shade-inhabiting Costa Rican rain forest butterfly (Hesperiidae). On both males and females, two types of whiteness occur: angle dependent (dull or bright) and angle independent, which differ in the microstructure, orientation, and associated properties of their scales. Some spots on the male wings are absent from the female wings. Whether the angle-dependent whiteness is bright or dull depends on the observation directions. The angle-dependent scales also show enhanced retro-reflection. We speculate that the biological functions and evolution of Carystoides spot patterns, scale structures, and their varying whiteness are adaptations to butterfly's low light habitat and to airflow experienced on the wing base vs. wing tip.

Light availability affects sex lability in a gynodioecious plant.

Sex lability (i.e., gender diphasy) in plants is classically linked to the larger resource needs associated with the female sexual function (i.e., seed production) compared to the male function (i.e., pollen production). Sex lability in response to the environment is extensively documented in dioecious species, but has been largely overlooked in gynodioecious plants. Here, we tested whether environmental conditions induce sex lability in the gynodioecious Geranium sylvaticum. We conducted a transplantation experiment in the field where plants with different sex expression were reciprocally transplanted between high light and low light habitats. We measured plants' reproductive output and sex expression over four years. Our results show that sex expression was labile over the study period. The light level at the destination habitat had a significant effect on sexual expression and reproductive output, because plants decreased their reproductive output when transplanted to the low light habitat. Transplantation origin did not affect any parameter measured. This study shows that sex expression in Geranium sylvaticum is labile and related to light availability. Sexually labile plants did not produce more seeds or pollen, and thus, there was no apparent fitness gain in sexually labile individuals. Sex lability in gynodioecious plants may be more common than previously believed because detection of sex lability necessitates data on the same individuals over time, which is rare in sexually dimorphic herbaceous plants.

Ontogenetic responses of four plant species to additive and interactive effects of land‐use history, canopy structure and herbivory

Summary The strength of interactions among species is often highly variable in space and time, and a major challenge in understanding context‐dependent effects of herbivores lies in disentangling habitat‐mediated from herbivore‐mediated effects on plant performance. We conducted a landscape‐scale experiment that manipulated light availability in woodlands with either a history of agricultural use or no history of agricultural use and coupled this with performance measurements of three life stages on four perennial herbaceous species exposed to varying levels of herbivory. We found that the context‐dependent effects of herbivory on plant performance changed as plants grew: juvenile plant survival was reduced by herbivores in low‐light habitats whereas biomass of adult plants was reduced by a more diverse insect fauna in high‐light environments. A history of agricultural land use also had negative effects on seedling establishment and adult performance, independent of herbivory. Synthesis. This work experimentally separates the habitat‐mediated effects on plant performance from the herbivore‐mediated effects on plant performance and highlights how context‐dependent interactions depend on plant ontogenetic stages.

Habitat heterogeneity reflected in mesophotic reef sediments

Modern reef sediments reflect the physical and chemical characteristics of the environment as well as the local reef fauna. Analysis of sedimentary reef facies can thus provide a powerful tool in interpreting ancient reef deposits. However, few studies have attempted to differentiate sedimentary facies in mesophotic coral ecosystems, low light habitats defined as residing 30–150m below sea level. The low-angle shelf mesophotic coral ecosystem south of the northern U.S. Virgin Islands (USVI) consists of reefs with different structural characteristics ideal for studying the relationship between habitat variability and sedimentary facies. Textural, compositional, and geochemical analyses of surface sediments were used to identify mesophotic reef subfacies associated with distinct benthic communities and structural habitats. Sediment grain composition and bulk geochemistry were found to broadly record the distribution and abundance of coral and macroalgae communities, foundational mesophotic reef benthic organisms. Overall, sediment composition was found to be a good indicator of specific reef environments in low-angle mesophotic reef habitats. Sedimentological analyses indicate that hydrodynamic forces do not transport a significant amount of allochthonous sediment or potentially harmful terrigenous material to USVI mesophotic reefs. Episodic, maximum current velocities prevented deposition of most silt-size grains and smaller, but biological processes were found to have a greater influence on subfacies partitioning than hydrodynamic processes. Results provide a new analog for studies of ancient mesophotic coral ecosystem geological history and document the relationship between mesophotic reef subfacies, structural complexity, and habitat heterogeneity. They also demonstrate how mesophotic reefs along the same shelf system do not always share similar sedimentary characteristics and thus record a diverse set of ecological and environmental conditions.

Restricted use of nitrate and a strong preference for ammonium reflects the nitrogen ecophysiology of a light-limited red alga.

Ammonium and nitrate are important sources of inorganic nitrogen for coastal primary producers. Nitrate has higher energy requirement for uptake and assimilation, compared with ammonium, suggesting that it might be a more efficient nitrogen source for slow-growing, light-limited macroalgae. To address this hypothesis, we examined the nitrogen ecophysiology of Anotrichium crinitum, a rhodophyte macroalgae common in low-light habitats in New Zealand. We measured seasonal changes in seawater nitrate and ammonium concentrations and the concentration of nitrate and ammonium stored internally by A.crinitum. We determined the maximal uptake rates of nitrate and ammonium seasonally and grew A.crinitum in the laboratory with these nitrogen sources under two ecologically relevant saturating light levels. Our results show that field-harvested A.crinitum has a high affinity for ammonium and although it will grow when supplied exclusively with nitrate, internal nitrate pools are low and it is unable to take up nitrate without several days of acclimation to saturating light. Our data predict that A.crinitum would be able to sustain growth with ammonium as the sole source of nitrogen, a strategy that would help it survive under low-light conditions that prevail in the field.

Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal, keen-eyed predator

High visual acuity allows parallel processing of distant environmental features, but only when photons are abundant enough. Diurnal tiger beetles (Carabidae: Cicindelinae) have acute vision for insects and visually pursue prey in open, flat habitats. Their fast running speed causes motion blur that degrades visual contrast, forces stop-and-go pursuit and potentially impairs obstacle detection. We demonstrate here that vision is insufficient for obstacle detection during running, and show instead that antennal touch is both necessary and sufficient for obstacle detection. While running, tiger beetle vision appears to be photon-limited in a way reminiscent of animals in low-light habitats. Such animals often acquire wide-field spatial information through mechanosensation mediated by longer, more mobile appendages. We show that a nocturnal tiger beetle species waves its antennae in elliptical patterns typical of poorly sighted insects. While antennae of diurnal species are also used for mechanosensation, they are rigidly held forward with the tips close to the substrate. This enables timely detection of path obstructions followed by an increase in body pitch to avoid collision. Our results demonstrate adaptive mechanosensory augmentation of blurred visual information during fast locomotion, and suggest that future studies may reveal non-visual sensory compensation in other fast-moving animals.

Integrated assessment of the direct and indirect effects of resource gradients on tree species recruitment

Understanding the dynamics of tree establishment is critical to assess forests' composition, management practices, and current responses to global change. We carried out a field seedling transplant experiment to assess not only the direct effects of resources influencing recruitment of four tree species, but also their indirect and combined effects. Our analysis integrated first growing season demographic data together with estimates of mycorrhizal fungal colonization and resource availability (light, soil moisture, and soil nitrogen). Only by considering both the direct and indirect effects of resources we were able to account for most of the variability observed during seedling recruitment. Contrary to expectations, increasing light levels were not always beneficial for recruitment even in low light habitats, and soil moisture availability benefited seedling growth but not survival. In addition, mycorrhizal fungal colonization was not always favored by high light levels or by increasing soil moisture. Seedling survival for all species was lower in plots with higher arbuscular mycorrhizal fungi, while the association with ectomycorrhizal fungi varied from beneficial to detrimental. When integrating the direct, indirect, and interactive effects of resource availability and mycorrhizal fungal colonization on tree recruitment dynamics we found that species responded in a nonlinear fashion to increasing resource levels, and we also identified thresholds, i.e., shifts in the direction of the response, along the resource gradient. Our integrated assessment considerably outperformed a null model where only direct effects of resources were accounted for. These results illustrate how the combination of direct, indirect, and combined effects of driving variables better represents the complexity of the processes determining tree species recruitment than simple resource availability mechanisms.

The symbiosis between the gastropod Strombus gigas and the dinoflagellate Symbiodinium: An ontogenic journey from mutualism to parasitism

Dinoflagellates in the genus Symbiodinium form mutualistic symbioses not only with corals and other cnidarians but also with sponges, bivalves, and gastropods, including the commercially important queen conch, Strombus gigas (Linnaeus 1758). The eggs of S. gigas are aposymbiotic, but acquisition of Symbiodinium at the larval stage significantly enhances survival and growth, demonstrating the mutualistic nature of this symbiosis at the larval stage. The objective of the present study was to investigate the symbiotic association between adult S. gigas and Symbiodinium. S. gigas were collected along the Mexican Caribbean coast. Symbiodinium did not occur in the muscle tissue or gonads, the latter supporting the lack of symbionts in S. gigas' eggs. Symbiodinium cells were morphologically intact in all of the anatomical structures in which they were found that included, in order of abundance, the digestive gland and other parts of the digestive tract, gills, in connective tissue surrounding the mantle and foot, as well as in the nephridium and proboscis. The symbionts underwent mitosis in the mantle, stomach and digestive gland. The healthy condition of the Symbiodinium, evident in their morphology and mitotic index, was surprising given the low light habitat in which they were found in adult S. gigas. The shell of S. gigas is a strong light attenuator; a maximum of 21% of incident radiation passed through the conch shell and decreased exponentially with increasing shell length to a low of 0.1%. The maximal photochemical efficiencies (Fv/Fm) of the symbionts were low, suggesting that their photosynthetic activity would be suboptimal in the reduced light environment in which they are found. The large population of Symbiodinium may be maintained by the symbionts acquiring nutrients heterotrophically from the host. Consequently, in S. gigas, a mutualistic symbiont at the larval life stage may confer a net cost at the adult life stage.

Implications of geometric plasticity for maximizing photosynthesis in branching corals

Reef-building corals are an example of plastic photosynthetic organisms that occupy environments of high spatiotemporal variations in incident irradiance. Many phototrophs use a range of photoacclimatory mechanisms to optimize light levels reaching the photosynthetic units within the cells. In this study, we set out to determine whether phenotypic plasticity in branching corals across light habitats optimizes potential light utilization and photosynthesis. In order to do this, we mapped incident light levels across coral surfaces in branching corals and measured the photosynthetic capacity across various within-colony surfaces. Based on the field data and modelled frequency distribution of within-colony surface light levels, our results show that branching corals are substantially self-shaded at both 5 and 18 m, and the modal light level for the within-colony surface is 50 mu mol photons m(-2) s(-1). Light profiles across different locations showed that the lowest attenuation at both depths was found on the inner surface of the outermost branches, while the most self-shading surface was on the bottom side of these branches. In contrast, vertically extended branches in the central part of the colony showed no differences between the sides of branches. The photosynthetic activity at these coral surfaces confirmed that the outermost branches had the greatest change in sun- and shade-adapted surfaces; the inner surfaces had a 50 % greater relative maximum electron transport rate compared to the outer side of the outermost branches. This was further confirmed by sensitivity analysis, showing that branch position was the most influential parameter in estimating whole-colony relative electron transport rate (rETR). As a whole, shallow colonies have double the photosynthetic capacity compared to deep colonies. In terms of phenotypic plasticity potentially optimizing photosynthetic capacity, we found that at 18 m, the present coral colony morphology increased the whole-colony rETR, while at 5 m, the colony morphology decreased potential light utilization and photosynthetic output. This result of potential energy acquisition being underutilized in shallow, highly lit waters due to the shallow type morphology present may represent a trade-off between optimizing light capture and reducing light damage, as this type morphology can perhaps decrease long-term costs of and effect of photoinhibition. This may be an important strategy as opposed to adopting a type morphology, which results in an overall higher energetic acquisition. Conversely, it could also be that maximizing light utilization and potential photosynthetic output is more important in low-light habitats for Acropora humilis.

Survival in low light: photosynthesis and growth of a red alga in relation to measured in situ irradiance.

Reduced light availability for benthic primary producers as a result of anthropogenic activities may be an important driver of change in coastal seas. However, our knowledge of the minimum light requirements for benthic macroalgae limits our understanding of how these changes may affect primary productivity and the functioning of coastal ecosystems. This knowledge gap is particularly acute in deeper water, where the impacts of increased light attenuation will be most severe. We examined the minimum light requirements of Anotrichium crinitum, which dominates near the maximum depth limit for macroalgae throughout New Zealand and Southern Australia, and is a functional analog of rhodophyte macroalgae in temperate low-light (deep-water) habitats throughout the world. These data show that A.crinitum is a shade-adapted seaweed with modest light requirements for the initiation of net photosynthesis (1.49-2.25μmol photons·m(-2) ·s(-1) ) and growth (0.12-0.19mol photons·m(-2) ·d(-1) ). A.crinitum maintains high photosynthetic efficiency and pigment content and a low C:N ratio throughout the year and can maintain biomass under sub-compensation (critical) light levels for at least 5d. Nevertheless, in situ photon flux is less than the minimum light requirement for A.crinitum on at least 103d per annum and is rarely sufficient to saturate growth. These findings reinforce the importance of understanding the physiological response of macroalgae at the extremes of environmental gradients and highlight the need to establish minimum thresholds that modification of the subtidal light environment should not cross.