Up-regulation of bundle sheath electron transport capacity under limiting light in C4Setaria viridis

C4 photosynthesis is a biochemical pathway that operates across mesophyll and bundle sheath (BS) cells to increase CO2 concentration at the site of CO2 fixation. C4 plants benefit from high irradiance but their efficiency decreases under shade causing a loss of productivity in crop canopies. We investigated shade acclimation responses of a model NADP-ME monocot Setaria viridis focussing on cell-specific electron transport capacity. Plants grown under low light (LL) maintained CO2 assimilation rates similar to high light plants but had an increased chlorophyll and light-harvesting-protein content, predominantly in BS cells. Photosystem II (PSII) protein abundance, oxygen-evolving activity and the PSII/PSI ratio all increased in LL BS cells indicating a higher capacity for linear electron flow. PSI, ATP synthase, Cytochrome b6f and the chloroplastic NAD(P) dehydrogenase complex, which constitute the BS cyclic electron flow machinery, were all upregulated in LL plants. A decline in PEP carboxylase activity in mesophyll cells and a consequent shortage of reducing power in BS chloroplasts was associated with the more oxidised redox state of the plastoquinone pool in LL plants and the formation of PSII - light-harvesting complex II supercomplexes with an increased oxygen evolution rate. Our results provide evidence of a redox regulation of the supramolecular composition of Photosystem II in BS cells in response to shading. This newly identified link contributes to understanding the regulation of PSII activity in C4 plants and will support strategies for crop improvement including the engineering of C4 photosynthesis into C3 plants.Significance statementThe efficiency of C4 photosynthesis decreases under low irradiance causing a loss of productivity in crop canopies. We investigate shade acclimation of a model NADP-ME monocot, analysing cell-specific protein expression and electron transport capacity. We propose a regulatory pathway controlling abundance and activity of Photosystem II in bundle sheath cells in response to irradiance.

Sex-specific structural and functional leaf traits and sun–shade acclimation in the dioecious tree Pistacia vera (Anacardiaceae)

In dioecious species, sex-related adaptive strategies, influenced by natural and sexual selection, allow each sex to meet the specific demands of reproduction. Differences in ecophysiological traits between males and females may rely on innate differences in secondary sex traits such as structural and functional leaf traits. We tested structural sexual leaf dimorphism in Pistacia vera L. and the intersexual differences in sun–shade acclimation processes expected from the different adaptive strategies of males and females. Fifteen structural and functional leaf traits were compared in 50-year-old trees between females with low fruit load and males under sun and shade conditions. Despite the low additional energy investment in reproduction in females, remarkable sex effects in leaf structure and function were observed. Male trees had smaller leaves with significantly lower total conducting petiole area (TCA) and higher stomatal density, water use efficiency and concentration of phenolic compounds; females had larger leaves with greater thickness, leaf mass per area, TCA and maximum photosynthetic capacity per area (Amax,a). The higher Amax,a and stomatal conductance of female leaves were associated with their ~20-fold higher TCA compared with male trees. Females seem to invest more in high xylem efficiency and rates of C gain; males invest more in defence-protection. Sun–shade plastic responses were sex- and trait-specific, but the plasticity assessment indicated that both sexes have evolved an almost equal degree of phenotypic plasticity that allows them to perform optimally under varying environmental conditions. However, the trait-specific differences indicate that each sex displays a different strategy of optimisation.

Plant carbon and water fluxes in tropical montane cloud forests

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.

From ecophysiology to phenomics: some implications of photoprotection and shade–sun acclimation in situ for dynamics of thylakoids in vitro

Half a century of research into the physiology and biochemistry of sun–shade acclimation in diverse plants has provided reality checks for contemporary understanding of thylakoid membrane dynamics. This paper reviews recent insights into photosynthetic efficiency and photoprotection from studies of two xanthophyll cycles in old shade leaves from the inner canopy of the tropical trees Inga sapindoides and Persea americana (avocado). It then presents new physiological data from avocado on the time frames of the slow coordinated photosynthetic development of sink leaves in sunlight and on the slow renovation of photosynthetic properties in old leaves during sun to shade and shade to sun acclimation. In so doing, it grapples with issues in vivo that seem relevant to our increasingly sophisticated understanding of Δ pH-dependent, xanthophyll-pigment-stabilized non-photochemical quenching in the antenna of PSII in thylakoid membranes in vitro .

ACCLIMATION OF PHOTOSYNTHESIS AND GROWTH OF BANANA (MUSA SP.) TO NATURAL SHADE IN THE HUMID TROPICS

SUMMARYGrowth and photosynthetic performance of banana (Musa sp.) grown in three levels of natural shade (33, 55 and 77% reduction in incoming radiation) were compared to an unshaded control treatment. Net CO2 assimilation rates generally decreased with increasing shade. Chlorophyll fluorescence revealed short-term dynamic photoinhibition under high light conditions but no evidence of sustained photoinhibitory damage to photosystem II. Dynamic photoinhibition decreased with increasing shade, with the greatest depression in the variable to maximal fluorescence ratio (Fv/Fm) occurring in unshaded plants during the middle of the day. Specific leaf area and leaf area ratio increased proportionately with increasing shade, whilst the chlorophyll a/b ratio decreased, reflecting a greater efficiency of light utilization under shady conditions. The optimum shade level for photosynthetic productivity would be one at which the level of photosynthetic photon flux density (PPFD) is high enough to saturate CO2 assimilation but low enough to induce shade acclimation and to reduce photoinhibition. Under the conditions studied here, the saturation level of PPFD was around 1000 μmol m−2 s−1, a light level typical of the tree-based intercropping systems in which banana is commonly grown in the tropics.

Coordinated variation in ecophysiological properties among life stages and tissue types in an invasive perennial forb of semiarid shrub steppe

Semiarid lands of North America are vulnerable to invasion by exotic forbs that frequently have an extended phenology compared with native herbaceous species. We hypothesized that photosynthetic stems, rosette leaves, and cauline leaves of Centaurea maculosa Lam. (knapweed) would differ in ecophysiological specialization and that these differences would enhance plant carbon uptake across seasonal variations in microclimate. Photosynthesis, water relations, and morphological features of C. maculosa were measured under natural variations in temperature and soil water, and under manipulated light environments, in adults compared with seedlings having only rosette leaves. Carbon gain was greater in adults than in seedlings owing to high photosynthesis in cauline and rosette leaves when water was abundant. Otherwise, photosynthesis was relatively lower but persistent through drought in all tissues and ages until senescence. Photosynthesis decreased with water stress in all tissues except stems. Stems comprised up to 36% of photosynthetic area following senescence of rosette leaves during seasonal drought. Seedlings expressed shade acclimation compared with rosette and especially cauline leaves of adults, although adult rosette leaves had flexible photosynthetic light responses. Contrasting ecophysiological specializations of photosynthetic tissues enhance carbon gain of adult C. maculosa as light, water, and temperature vary during its relatively long growth season.