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Solar irradiance is the source of exergy for all living organisms. Photosynthesis in primeval organisms generates the food for the other species. It also provides the chemical energy in biomass that is used as fuel. Energy conversions in humans produce mechanical work using food as the exergy source. The food intake; the metabolic and thermic processes in the human body; the production of adenosine triphosphate (ATP); and the conversion of the ATP energy into mechanical work are analyzed using the principles of thermodynamics. An interesting conclusion is that humans have evolved as inefficient energy conversion systems, with food-to-work exergetic efficiencies close to 10%. The analyses and a number of examples in this chapter elucidate the application of thermodynamics to biological processes including: production and use of biomass; exergy value of nutrients; exergy spent for vital processes, such as respiration, blood circulation, and maintenance of body temperature; and exergy spent in sports, such as weight-lifting, walking races, the marathon, and bicycling. The chapter also surveys the relationship between exergy destruction, the state of health, aging, and life expectancy.
Here, I outline the various sources of energy that different life-forms use. The most fundamental division of life from an energy perspective is that between self-feeders that can utilize non-biological forms of energy such as light (autotrophs) and those that feed on other organisms (heterotrophs). Further subdivision of the autotroph category shows that not all of these organisms conduct what can be called ordinary photosynthesis – the type that yields oxygen. Some autotrophs conduct non-oxygenic photosynthesis, while others do not use light at all, but rather utilize chemical energy of various kinds in processes that are collectively called chemosynthesis. Next, I consider the flow of energy in ecosystems and note the limitations in the efficiency of energy transfer between trophic levels, but also the limitations of the trophic-level concept itself. Finally, I note that the biosphere is a realm of decreasing entropy: processes that contribute to this decrease include evolution, embryological development, and ecological succession. The decreasing entropy of the biosphere is perfectly compatible with the second law of thermodynamics as this law only applies to closed systems.
This study presents two years of characterization of a warm temperate rhodolith bed in order to analyse how certain environmental changes influence the community ecology. The biomass of rhodoliths and associated species were analysed during this period and in situ experiments were conducted to evaluate the primary production, calcification and respiration of the dominant species of rhodoliths and epiphytes. The highest total biomass of rhodoliths occurred during austral winter. Lithothamnion crispatum was the most abundant rhodolith species in austral summer. Epiphytic macroalgae occurred only in January 2015, with Padina gymnospora being the most abundant. Considering associated fauna, the biomass of Mollusca increased from February 2015 to February 2016. Population densities of key reef fish species inside and around the rhodolith beds showed significant variations in time. The densities of grouper (carnivores/piscivores) increased in time, especially from 2015 to 2016. On the other hand, grunts (macroinvertebrate feeders) had a modest decrease over time (from 2014 to 2016). Other parameters such as primary production and calcification of L. crispatum were higher under enhanced irradiance, yet decreased in the presence of P. gymnospora. Community structure and physiological responses can be explained by the interaction of abiotic and biotic factors, which are driven by environmental changes over time. Biomass changes can indicate that herbivores play a role in limiting the growth of epiphytes, and this is beneficial to the rhodoliths because it decreases competition for environmental resources with fleshy algae.
The Kepler data show that habitable small planets orbiting Red Dwarf stars (RDs) are abundant, and hence might be promising targets to look at for biomarkers and life. Planets orbiting within the habitable zone of RDs are close enough to be tidally locked. Some recent works have cast doubt on the ability of planets orbiting RDs to support life. In contrast, it is shown that temperatures suitable for liquid water and even for organic molecules may exist on tidally locked planets (TLPs) of RDs for a wide range of atmospheres. We chart the surface temperature distribution as a function of the irradiation, greenhouse factor and heat circulation. The habitability boundaries and their dependence on the atmospheric properties are derived. By extending our previous analyses of TLPs, we find that tidally locked as well as synchronous (not completely locked) planets of RDs and K-type stars may support life, for a wider range of orbital distance and atmospheric conditions than previously thought. In particular, it is argued that life clement environments may be possible on tidally locked and synchronously orbiting planets of RDs and K-type stars, with conditions supporting oxygenic photosynthesis, which on Earth was a key to complex life. Different climate projections and the biological significance of tidal locking on putative complex life are reviewed. We show that when the effect of continuous radiation is taken into account, the photo-synthetically active radiation available on TLPs, even of RDs, could produce a high-potential plant productivity, in analogy to mid-summer growth at high latitudes on Earth. Awaiting the findings of TESS and JWST, we discuss the implications of the above arguments to the detection of biomarkers such as liquid water and oxygen, as well as to the abundance of biotic planets and life.
Photosynthesis offers a convenient means of sustaining biospheres. We quantify the constraints for photosynthesis to be functional on the permanent nightside of tidally locked rocky exoplanets via reflected light from their exomoons. We show that the exomoons must be at least half the size of Earth's moon in order for conventional oxygenic photosynthesis to operate. This scenario of photosynthesis is unlikely for exoplanets around late-type M-dwarfs due to the low likelihood of large exomoons and their orbital instability over long timescales. Subsequently, we investigate the prospects for photosynthesis on habitable exomoons via reflected light from the giant planets that they orbit. Our analysis indicates that such photosynthetic biospheres are potentially sustainable on these moons except those around late-type M-dwarfs. We conclude our analysis by delineating certain physiological and biochemical features of photosynthesis and other carbon fixation pathways, and the likelihood of their evolution on habitable planets and moons.
The physiology of mesophotic Scleractinia varies with depth in response to environmental change. Previous research has documented trends in heterotrophy and photosynthesis with depth, but has not addressed between-site variation for a single species. Environmental differences between sites at a local scale and heterogeneous microhabitats, because of irradiance and food availability, are likely important factors when explaining the occurrence and physiology of Scleractinia. Here, 108 colonies of Agaricia lamarcki were sampled from two locations off the coast of Utila, Honduras, distributed evenly down the observed 50 m depth range of the species. We found that depth alone was not sufficient to fully explain physiological variation. Pulse Amplitude-Modulation fluorometry and stable isotope analyses revealed that trends in photochemical and heterotrophic activity with depth varied markedly between sites. Our isotope analyses do not support an obligate link between photosynthetic activity and heterotrophic subsidy with increasing depth. We found that A. lamarcki colonies at the bottom of the species depth range can be physiologically similar to those nearer the surface. As a potential explanation, we hypothesize sites with high topographical complexity, and therefore varied microhabitats, may provide more physiological niches distributed across a larger depth range. Varied microhabitats with depth may reduce the dominance of depth as a physiological determinant. Thus, A. lamarcki may ‘avoid’ changes in environment with depth, by instead existing in a subset of favourable niches. Our observations correlate with site-specific depth ranges, advocating for linking physiology and abiotic profiles when defining the distribution of mesophotic taxa.
Cyanobacteria and plants carry out oxygenic photosynthesis. They use water to generate the atmospheric oxygen we breathe and carbon dioxide to produce the biomass serving as food, feed, fibre and fuel. This paper scans the emergence of structural and mechanistic understanding of oxygen evolution over the past 50 years. It reviews speculative concepts and the stepped insight provided by novel experimental and theoretical techniques. Driven by sunlight photosystem II oxidizes the catalyst of water oxidation, a hetero-metallic Mn4CaO5(H2O)4 cluster. Mn3Ca are arranged in cubanoid and one Mn dangles out. By accumulation of four oxidizing equivalents before initiating dioxygen formation it matches the four-electron chemistry from water to dioxygen to the one-electron chemistry of the photo-sensitizer. Potentially harmful intermediates are thereby occluded in space and time. Kinetic signatures of the catalytic cluster and its partners in the photo-reaction centre have been resolved, in the frequency domain ranging from acoustic waves via infra-red to X-ray radiation, and in the time domain from nano- to milli-seconds. X-ray structures to a resolution of 1.9 Å are available. Even time resolved X-ray structures have been obtained by clocking the reaction cycle by flashes of light and diffraction with femtosecond X-ray pulses. The terminal reaction cascade from two molecules of water to dioxygen involves the transfer of four electrons, two protons, one dioxygen and one water. A rigorous mechanistic analysis is challenging because of the kinetic enslaving at millisecond duration of six partial reactions (4e−, 1H+, 1O2). For the time being a peroxide-intermediate in the reaction cascade to dioxygen has been in focus, both experimentally and by quantum chemistry. Homo sapiens has relied on burning the products of oxygenic photosynthesis, recent and fossil. Mankind's total energy consumption amounts to almost one-fourth of the global photosynthetic productivity. If the average power consumption equalled one of those nations with the highest consumption per capita it was four times greater and matched the total productivity. It is obvious that biomass should be harvested for food, feed, fibre and platform chemicals rather than for fuel.
Lichens are one of the common dominant biota in biological soil crusts (biocrusts), a community that is one of the largest in extent in the world. Here we present a summary of the main features of the lifestyle of soil crust lichens, emphasizing their habitat, ecophysiology and versatility. The soil crust is exposed to full light, often to high temperatures and has an additional water source, the soil beneath the lichens. However, despite the open nature of the habitat the lichens are active under shady and cooler conditions and avoid climate extremes of high temperature and light. In temperate and alpine habitats they can also be active for long periods, several months in some cases. They show a mixture of physiological constancy (e.g. similar activity periods and net photosynthetic rates) but also adaptations to the habitat (e.g. the response of net photosynthesis to thallus water content can differ for the same lichen species in Europe and the USA and some species show extensive rhizomorph development). Despite recent increased research, aspects of soil crust ecology, for example under snow, remain little understood.
Chilling injury is an important natural stress that can threaten cotton production, especially at the sowing and seedling stages in early spring. It is therefore important for cotton production to improve chilling tolerance at these stages. The current work examines the potential for glycine betaine (GB) treatment of seeds to increase the chilling tolerance of cotton at the seedling stage. Germination under cold stress was increased significantly by GB treatment. Under low temperature, the leaves of seedlings from treated seeds exhibited a higher net photosynthetic rate (PN), higher antioxidant enzyme activity including superoxide dismutase, ascorbate peroxidase and catalase, lower hydrogen peroxide (H2O2) content and less damage to the cell membrane. Enzyme activity was correlated negatively with H2O2 content and degree of damage to the cell membrane but correlated positively with GB content. The experimental results suggested that although GB was only used to treat cotton seed, the beneficial effect caused by the preliminary treatment of GB could play a significant role during germination that persisted to at least the four-leaf seedling stage. Therefore, it is crucial that this method is employed in agricultural production to improve chilling resistance in the seedling stage by soaking the seeds in GB.
The citrus leafminer (CLM), Phyllocnistis citrella Stainton causes injury to citrus and related species in the Rutaceae family. The damage that the CLM larvae can cause is significant in citrus plantations. We tested two citrus cultivars — ‘Kinnow’ (Citrus reticulata Blanco) and ‘Fairchild’ (a hybrid of Citrus reticulata Clementine x Orlando Tangelo) — to quantify CLM larvae infestation and effect on the physiology of the citrus cultivars. We then compared the CLM larval weight with its associated damage. To calculate infestation level, mine area and total leaf area, we used the image analysis technique. The infestation level of CLM was higher in ‘Fairchild’ than in ‘Kinnow’ cultivar of citrus. For both cultivars, larval weight of CLM was directly proportional to the amount of mines generated. Taken together, the results of this study suggest that the mines that CLM larvae generate pose significant effect on the net photosynthetic rates and water use efficiency of citrus nursery plants. These results will help improve our understanding of the interaction between CLM and citrus nursery plants and effect of the pest on the yield potential of the crop.
Hemiparasitic plants obtain water and solutes from their hosts, but much remains to be learned about these transfers. We used a forest girdling experiment to investigate how leaf gas exchange, carbon and nitrogen cycling in the root hemiparasite Melampyrum lineare Desr. responded to disturbance and changes in physiology of potential host trees. By preventing belowground C allocation by 35% of the canopy, girdling decreased the starch and soluble sugar contents of bulk forest floor fine roots. Photosynthetic rates of M. lineare were statistically significantly lower in the girdled plot, but their hypothesized drivers (foliar N, stomatal conductance and transpiration) had no statistically significant differences between girdled and non-girdled plots. However, M. lineare in the girdled plot had higher foliar C concentrations and Δ14C than in the control plot, suggesting possible photosynthetic down-regulation in the girdled plot due to influx of older (e.g., host-derived) C into the leaves of M. lineare. Within the girdled plot (but not the control plot), M. lineare foliar C concentrations were positively correlated with foliar Δ14C and δ15N, suggesting that M. lineare may respond to changes in both C and N biogeochemistry during the decline of dominant canopy species.
Within Antarctica there are large gradients both in climate and in vegetation which offer opportunities to investigate links between the two. The activity (% total time active) of lichens and bryophytes in hydric and xeric environments was monitored at Livingston Island (62°39'S). This adds a northern site with a maritime, cloudy climate to previous studies in the southern Antarctic Peninsula and the Dry Valleys (78°S). Annual activity increases northwards from less than 1% to nearly 100%. There is a major and consistent difference between hydric sites which, with snow melt, can be 100% active in summer months even in the Dry Valleys, and xeric sites which, depending on precipitation, rarely exceed 80% activity even at Livingston Island. Mosses dominate hydric sites and lichens the xeric sites all along the gradient. Mean temperatures when active are 2–4°C at all sites, as liquid water is required. Light is a potential major stress reaching 880 µmol m-2 s-1 when active in continental sites. The lack of extremes in temperatures and light combined with high activity levels means that summer at Livingston Island is one of the better sites for lichen and bryophyte growth in the world.
Tomato grafting is practiced worldwide as an innovative approach to manage stress from drought, waterlogging, insects, and diseases. Metribuzin is a commonly used herbicide in tomato but has potential to cause injury after application if plants are under stress. The influence of metribuzin on grafted tomato under drought-stress has not been studied. Greenhouse experiments were conducted in Raleigh, NC to determine the tolerance of drought-stressed grafted and non-grafted tomato to metribuzin. The tomato cultivar ‘Amelia’ was used as the scion in grafted tomato, and for the non-grafted control. Two hybrid tomato ‘Beaufort’ and ‘Maxifort’ were used as rootstocks for grafted plants. Drought-stress treatments included: no drought-stress; 3 d of drought-stress before metribuzin application with no drought-stress after application (3 d DSB); and 3 d of drought-stress before metribuzin application with 3 d of drought-stress after application (3 d DSBA). Metribuzin was applied at 550 g ai ha−1. No difference in injury from metribuzin was observed in grafted and non-grafted plants. However, at 7 and 14 d after metribuzin treatment (DMT), less injury was observed on tomato in the 3 d DSBA treatment (5 and 2% injury, respectively) than on plants in the 3 d DSB treatment (15 and 8% injury, respectively) or those that were never drought-stressed (18 and 11% injury, respectively). Photosynthesis and stomatal conductance measured prior to metribuzin application were reduced similarly in grafted and non-grafted tomato subjected to drought-stress. Photosynthesis and stomatal conductance of grafted and non-grafted tomato at 7 DMT was not different among drought-stress treatments or metribuzin treatments. Grafted and non-grafted tomato plants under drought-stress exhibit similar tolerance to metribuzin. The risk of metribuzin injury to grafted tomato under drought-stress is similar to non-grafted tomato.
A greater relative allocation of phosphorus (P) to photosynthetically active cells functions to maintain a rapid photosynthesis under P limitation, and may be a key mechanism of plants to use P efficiently. This mechanism has not been studied in tropical trees despite the productivity of tropical forests often being limited by P. In this study, the spatial distribution of P among tissues on a cross-section of leaf lamina was analysed for 13 tree species from P-limited sites on Mount Kinabalu, Borneo. Most species showed greater P concentration in palisade mesophyll than in spongy mesophyll and epidermal tissues, suggesting that tropical trees under P limitation localize foliar P in photosynthetic palisade mesophyll.
pCO2/pH perturbation experiments were carried out under two different pCO2 levels to evaluate effects of CO2-driven ocean acidification on semi-continuous cultures of the marine diatom Skeletonema pseudocostatum CSA48. Under higher pCO2/lowered pH conditions, our results showed that CO2-driven acidification had no significant impact on growth rate, chlorophyll-a, cellular abundance, gross photosynthesis, dark respiration, particulate organic carbon and particulate organic nitrogen between CO2-treatments, suggesting that S. pseudocostatum is adapted to tolerate changes of ~0.5 units of pH under high pCO2 conditions. However, dissolved organic carbon (DOC) concentration and DOC/POC ratio were significantly higher at high pCO2, indicating that a greater partitioning of organic carbon into the DOC pool was stimulated by high CO2/low pH conditions. Total fatty acids (FAs) were significantly higher under low pCO2 conditions. The composition of FAs changed from low to high pCO2, with an increase in the concentration of saturated and a reduction of monounsaturated FAs. Polyunsaturated FAs did not show significant differences between pCO2 treatments. Our results lead to the conclusion that the balance between negative or null effect on S. pseudocostatum ecophysiology upon low pH/high pCO2 conditions constitute an important factor to be considered in order to evaluate the global effect of rising atmospheric CO2 on primary productivity in coastal ocean. We found a significant decrease in total FAs, however no indications were found for a detrimental effect of ocean acidification on the nutritional quality in terms of essential fatty acids.
Tropical montane cloud forests (TMCFs) are dynamic ecosystems defined by frequent, but intermittent, contact with fog. The resultant microclimate can vary considerably over short spatial and temporal scales, affecting the ecophysiology of TMCF plants. We synthesized research to date on TMCF carbon and water fluxes at the scale of the leaf, plant and ecosystem and then contextualized this synthesis with tropical lowland forest ecosystems. Mean light-saturated photosynthesis was lower than that of lowland forests, probably due to the effects of persistent reduced radiation leading to shade acclimation. Scaled to the ecosystem, measures of annual net primary productivity were also lower. Mean rates of transpiration, from the scale of the leaf to the ecosystem, were also lower than in lowland sites, likely due to lower atmospheric water demand, although there was considerable overlap in range. Lastly, although carbon use efficiency appears relatively invariant, limited evidence indicates that water use efficiency generally increases with altitude, perhaps due to increased cloudiness exerting a stronger effect on vapour pressure deficit than photosynthesis. The results reveal clear differences in carbon and water balance between TMCFs and their lowland counterparts and suggest many outstanding questions for understanding TMCF ecophysiology now and in the future.
Several monodominant rain-forest trees in New Caledonia have population size structures suggesting establishment following large-scale disturbance, with eventual replacement by shade-tolerant species predicted in the absence of future disturbance. Links of dominance and population dynamics to leaf-level photosynthesis were investigated in seedlings of 20 tree species from these forests, grown in experimental sun and shade conditions. In particular, we tested whether episodically regenerating (ER) species, including monodominants, have higher assimilation rates at high irradiances and lower tolerance of shade than continuously regenerating species (CR). ER species had higher maximum net assimilation rates (Amax-area) in sun plants (9.6 ± 0.4 μmol m−2 s−1) than CR species (6.2 ± 0.3 μmol m−2 s−1) and high plasticity, typical of shade-intolerant species, but monodominant species did not differ from other ER species. CR species had leaf-level traits consistent with shade tolerance, including lower dark respiration rates (Rd-area = 0.47 ± 0.03μmol m−2 s−1; Rd-mass = 7 ± 1 nmol g−1 s−1) than ER species (Rd-area = 0.63 ± 0.06 μmol m−2 s−1; Rd-mass = 11 ± 2 nmol g−1 s−1) in shade plants. Hence leaf-level assimilation traits were largely consistent with regeneration patterns, but do not explain why some shade-intolerant species can achieve monodominance.
A ‘habitable zone’ of a star is defined as a range of orbits within which a rocky planet can support liquid water on its surface. The most intriguing question driving the search for habitable planets is whether they host life. But is the age of the planet important for its habitability? If we define habitability as the ability of a planet to beget life, then probably it is not. After all, life on Earth has developed within only ~800 Myr after its formation – the carbon isotope change detected in the oldest rocks indicates the existence of already active life at least 3.8 Gyr ago. If, however, we define habitability as our ability to detect life on the surface of exoplanets, then age becomes a crucial parameter. Only after life had evolved sufficiently complex to change its environment on a planetary scale, can we detect it remotely through its imprint on the atmosphere – the so-called biosignatures, out of which the photosynthetic oxygen is the most prominent indicator of developed (complex) life as we know it. Thus, photosynthesis is a powerful biogenic engine that is known to have changed our planet's global atmospheric properties. The importance of planetary age for the detectability of life as we know it follows from the fact that this primary process, photosynthesis, is endothermic with an activation energy higher than temperatures in habitable zones, and is sensitive to the particular thermal conditions of the planet. Therefore, the onset of photosynthesis on planets in habitable zones may take much longer time than the planetary age. The knowledge of the age of a planet is necessary for developing a strategy to search for exoplanets carrying complex (developed) life – many confirmed potentially habitable planets are too young (orbiting Population I stars) and may not have had enough time to develop and/or sustain detectable life. In the last decade, many planets orbiting old (9–13 Gyr) metal-poor Population II stars have been discovered. Such planets had had enough time to develop necessary chains of chemical reactions and may carry detectable life if located in a habitable zone. These old planets should be primary targets in search for the extraterrestrial life.
Genetic inter-communication of the nucleic-organellar dual in eukaryotes is dominated by DNA-directed phenomena. RNA regulatory circuits have also been observed in artificial laboratory prototypes where gene transfer events are reconstructed, but they are excluded from the primary norm due to their rarity. Recent technical advances in organellar biotechnology, genome engineering and single-molecule tracking give novel experimental insights on RNA metabolism not only at cellular level, but also on organismal survival. Here, I put forward a hypothesis for RNA's involvement in gene piece transfer, taken together the current knowledge on the primitive RNA character as a biochemical modulator with model organisms from peculiar natural habitats. It is proposed that RNA molecules of special structural signature and functional identity can drive evolution, integrating the ecological pressure of environmental oscillations into genome imprinting by buffering-out epigenetic aberrancies.