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Tree growth is generally considered to be temperature limited at upper elevation treelines, yet climate factors controlling tree growth at semiarid treelines are poorly understood. We explored the influence of climate on stem growth and stable isotopes for Polylepis tarapacana Philipi, the world's highest elevation tree species, which is found only in the South American Altiplano. We developed tree-ring width index (RWI), oxygen (δ18O) and carbon (δ13C) chronologies for the last 60 years at four P. tarapacana stands located above 4400 m in elevation, along a 500 km latitude aridity gradient. Total annual precipitation decreased from 300 to 200 mm from the northern to the southern sites. We used RWI as a proxy of wood formation (carbon sink) and isotopic tree-ring signatures as proxies of leaf-level gas exchange processes (carbon source). We found distinct climatic conditions regulating carbon sink processes along the gradient. Current growing-season temperature regulated RWI at northern-wetter sites, while prior growing-season precipitation determined RWI at arid southern sites. This suggests that the relative importance of temperature to precipitation in regulating tree growth is driven by site water availability. By contrast, warm and dry growing seasons resulted in enriched tree-ring δ13C and δ18O at all study sites, suggesting that similar climate conditions control carbon-source processes along the gradient. Site-level δ13C and δ18O chronologies were significantly and positively related at all sites, with the strongest relationships among the southern drier stands. This indicates an overall regulation of intercellular carbon dioxide via stomatal conductance for the entire P. tarapacana network, with greater stomatal control when aridity increases. This manuscript also highlights a coupling (decoupling) between physiological processes at leaf level and wood formation as a function of similarities (differences) in their climatic sensitivity. This study contributes to a better understanding and prediction of the response of high-elevation Polylepis woodlands to rapid climate changes and projected drying in the Altiplano., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

The mangrove Avicennia marina adjusts internal salt concentrations by foliar salt secretion. Deliquescence of accumulated salt causes leaf wetting that may provide a water source for salt-secreting plants in arid coastal wetlands where high nocturnal humidity can usually support deliquescence whereas rainfall events are rare. We tested the hypotheses that salt deliquescence on leaf surfaces can drive top-down rehydration, and that such absorption of moisture from unsaturated atmospheres makes a functional contribution to dry season shoot water balances. Sap flow and water relations were monitored to assess the uptake of atmospheric water by branches during shoot wetting events under natural and manipulated microclimatic conditions. Reverse sap flow rates increased with increasing relative humidity from 70% to 89%, consistent with function of salt deliquescence in harvesting moisture from unsaturated atmospheres. Top-down rehydration elevated branch water potentials above those possible from root water uptake, subsidising transpiration rates and reducing branch vulnerability to hydraulic failure in the subsequent photoperiod. Absorption of atmospheric moisture harvested through deliquescence of salt on leaf surfaces enhances water balances of Avicennia marina growing in hypersaline wetlands under arid climatic conditions. Top-down rehydration from these frequent, low intensity wetting events contributes to prevention of carbon starvation and hydraulic failure during drought., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

Until recently, few data were available on photosynthesis and its underlying mechanistically limiting factors in plants, other than crops and model species. Currently, a new large pool of data from extant representatives of basal terrestrial plant groups is emerging, allowing exploration of how photosynthetic capacity (A max ) increases from minimum values in bryophytes to maximum in tracheophytes, which is associated to an optimization of the balance between its limiting factors. From predominant mesophyll conductance limitation (l m ) in bryophytes and lycophytes (fern allies) to stomatal conductance (l s ) and l m colimitation in pteridophytes (ferns) and gymnosperms, a balanced colimitation by the three limitations is finally reached in angiosperms. We discuss the implications of this new knowledge for future biotechnological attempts to improve crop photosynthesis., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Many clonal plants produce vegetative recruits that remain connected to the parent plant. Such connections permit resource sharing among ramets, explaining the high survival rates of vegetative recruits during establishment under suboptimal conditions for sexual regeneration. We propose that differences in the regeneration niches of sexual and vegetative recruits reflect different physiological adjustments caused by parental supply of resources to the ramets. We conducted ecophysiological measurements in saplings and root suckers of Eucryphia cordifolia Cav., a tree species of the temperate rainforest of southern South America. We compared the following traits of saplings and suckers: gas exchange at the leaf level, crown architecture, daily crown carbon balance, biomass allocation to above-ground tissues (leaf-to-stem mass ratio, leaf mass area and leaf area ratio), xylem anatomy traits (lumen vessel fraction, vessel density and size) and stem ring width. We also correlated the growth rates of saplings and suckers with relevant environmental data (light and climate). Saplings showed morphological, architectural and physiological traits that enhance daily crown carbon balance and increase water-use efficiency, in order to supply their growth demands while minimizing water loss per unit of carbon gained. The radial growth of saplings diminished under dry conditions, which suggests a strong stomatal sensitivity to water availability. Suckers have low stomatal conductance, likely because the carbon supplied by the parent plant diminishes the necessity of high rates of photosynthesis. The low responsiveness of sucker growth to temporal changes in water availability also supports the existence of parental supply. The physiological differences between sexual and vegetative recruits satisfactorily explain the ecological niche of E. cordifolia, with saplings restricted to more closed and humid sites., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Water limitation is a major global constraint for plant productivity that is likely to be exacerbated by climate change. Hence, improving plant water use efficiency (WUE) has become a major goal for the near future. At the leaf level, WUE is the ratio between photosynthesis and transpiration. Maintaining high photosynthesis under water stress, while improving WUE requires either increasing mesophyll conductance (gm ) and/or improving the biochemical capacity for CO2 assimilation-in which Rubisco properties play a key role, especially in C3 plants at current atmospheric CO2 . The goals of the present analysis are: (1) to summarize the evidence that improving gm and/or Rubisco can result in increased WUE; (2) to review the degree of success of early attempts to genetically manipulate gm or Rubisco; (3) to analyse how gm , gsw and the Rubisco's maximum velocity (Vcmax ) co-vary across different plant species in well-watered and drought-stressed conditions; (4) to examine how these variations cause differences in WUE and what is the overall extent of variation in individual determinants of WUE; and finally, (5) to use simulation analysis to provide a theoretical framework for the possible control of WUE by gm and Rubisco catalytic constants vis-à-vis gsw under water limitations., (© 2015 John Wiley & Sons Ltd.)

Ferns and fern allies have low photosynthetic rates compared with seed plants. Their photosynthesis is thought to be limited principally by physical CO2 diffusion from the atmosphere to chloroplasts. The aim of this study was to understand the reasons for low photosynthesis in species of ferns and fern allies (Lycopodiopsida and Polypodiopsida). We performed a comprehensive assessment of the foliar gas-exchange and mesophyll structural traits involved in photosynthetic function for 35 species of ferns and fern allies. Additionally, the leaf economics spectrum (the interrelationships between photosynthetic capacity and leaf/frond traits such as leaf dry mass per unit area or nitrogen content) was tested. Low mesophyll conductance to CO2 was the main cause for low photosynthesis in ferns and fern allies, which, in turn, was associated with thick cell walls and reduced chloroplast distribution towards intercellular mesophyll air spaces. Generally, the leaf economics spectrum in ferns follows a trend similar to that in seed plants. Nevertheless, ferns and allies had less nitrogen per unit DW than seed plants (i.e. the same slope but a different intercept) and lower photosynthesis rates per leaf mass area and per unit of nitrogen., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Plant growth at extremely high elevations is constrained by high daily thermal amplitude, strong solar radiation and water scarcity. These conditions are particularly harsh in the tropics, where the highest elevation treelines occur. In this environment, the maintenance of a positive carbon balance involves protecting the photosynthetic apparatus and taking advantage of any climatically favourable periods. To characterize photoprotective mechanisms at such high elevations, and particularly to address the question of whether these mechanisms are the same as those previously described in woody plants along extratropical treelines, we have studied photosynthetic responses in Polylepis tarapacana Philippi in the central Andes (18°S) along an elevational gradient from 4300 to 4900 m. For comparative purposes, this gradient has been complemented with a lower elevation site (3700 m) where another Polylepis species (P. rugulosa Bitter) occurs. During the daily cycle, two periods of photosynthetic activity were observed: one during the morning when, despite low temperatures, assimilation was high; and the second starting at noon when the stomata closed because of a rise in the vapour pressure deficit and thermal dissipation is prevalent over photosynthesis. From dawn to noon there was a decrease in the content of antenna pigments (chlorophyll b and neoxanthin), together with an increase in the content of xanthophyll cycle carotenoids. These results could be caused by a reduction in the antenna size along with an increase in photoprotection. Additionally, photoprotection was enhanced by a partial overnight retention of de-epoxized xanthophylls. The unique combination of all of these mechanisms made possible the efficient use of the favourable conditions during the morning while still providing enough protection for the rest of the day. This strategy differs completely from that of extratropical mountain trees, which uncouple light-harvesting and energy-use during long periods of unfavourable, winter conditions., (Published by Oxford University Press on behalf of the Annals of Botany Company.)

Ferns are thought to have lower photosynthetic rates than angiosperms and they lack fine stomatal regulation. However, no study has directly compared photosynthesis in plants of both groups grown under optimal conditions in a common environment. We present a common garden comparison of seven angiosperms and seven ferns paired by habitat preference, with the aims of (1) confirming that ferns do have lower photosynthesis capacity than angiosperms and quantifying these differences; (2) determining the importance of diffusional versus biochemical limitations; and (3) analysing the potential implication of leaf anatomical traits in setting the photosynthesis capacity in both groups. On average, the photosynthetic rate of ferns was about half that of angiosperms, and they exhibited lower stomatal and mesophyll conductance to CO2 (gm ), maximum velocity of carboxylation and electron transport rate. A quantitative limitation analysis revealed that stomatal and mesophyll conductances were co-responsible for the lower photosynthesis of ferns as compared with angiosperms. However, gm alone was the most constraining factor for photosynthesis in ferns. Consistently, leaf anatomy showed important differences between angiosperms and ferns, especially in cell wall thickness and the surface of chloroplasts exposed to intercellular air spaces., (© 2014 John Wiley & Sons Ltd.)

Eucryphia cordifolia Cav. is a long-lived evergreen tree species, commonly found as a canopy emergent tree in the Chilean temperate rain forest. This species displays successive leaf cohorts throughout the entire growing season. Thus, full leaf expansion occurs under different environmental conditions during growing such as air temperature, vapor pressure deficit and the progress of moderate water stress (WS). These climate variations can be reflected as differences in anatomical and physiological characteristics among leaf cohorts. Thus, we investigated the potential adaptive role of different co-existing leaf cohorts in seedlings grown under shade, drought stress or a combination of the two. Photosynthetic and anatomical traits were measured in the first displayed leaf cohort and in a subsequent leaf cohort generated during the mid-season. Although most anatomical and photosynthetic pigments did not vary between cohorts, photosynthetic acclimation did occur in the leaf cohort and was mainly driven by biochemical processes such as leaf nitrogen content, Rubisco carboxylation capacity and maximal Photosystem II electron transport rather than CO2 diffusion conductance. Cohort acclimation could be relevant in the context of climate change, as this temperate rainforest will likely face some degree of summer WS even under low light conditions. We suggest that the acclimation of the photosynthetic capacity among current leaf cohorts represents a well-tuned mechanism helping E. cordifolia seedlings to face a single stress like shade or drought stress, but is insufficient to cope with simultaneous stresses., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The climate change conditions predicted for the end of the current century are expected to have an impact on the performance of plants under natural conditions. The variables which are foreseen to have a larger effect are increased CO2 concentration and temperature. Although it is generally considered CO2 assimilation rate could be increased by the increasing levels of CO2, it has been reported in previous studies that acclimation to high CO2 results in reductions of physiological parameters involved in photosynthesis, like the maximum carboxylation rate (Vc,max), stomatal conductance (gs) and mesophyll conductance to CO2 (gm). On the one hand, most of the previous modeling efforts have neglected the potential role played by the acclimation of gm to high CO2 and temperature. On the other hand, the effect of climate change on plant clades other than angiosperms, like ferns, has received little attention, and there are no studies evaluating the potential impact of increasing CO2 and temperature on these species. In this study we predicted responses of several representative species among angiosperms, gymnosperms and ferns to increasing CO2 and temperature. Our results show that species with lower photosynthetic capacity - such as some ferns and gymnosperms - would be proportionally more favored under these foreseen environmental conditions. The main reason for this difference is the lower diffusion limitation imposed by gs and gm in plants having high capacity for photosynthesis among the angiosperms, which reduces the positive effect of increasing CO2. However, this apparent advantage of low-diffusion species would be canceled if the two conductances - gs and gm - acclimate and are down regulated to high CO2, which is basically unknown, especially for gymnosperms and ferns. Hence, for a better understanding of different plant responses to future climate, studies are urged in which the actual photosynthetic response/acclimation to increased CO2 and temperature of ferns, gymnosperms and other under-evaluated plant groups is assessed., (Copyright © 2014 Elsevier Ireland Ltd. All rights reserved.)

Maximum photosynthesis rates in ferns are generally lower than those of seed plants, but little is known about the limiting factors, which are crucial to understand the evolution of photosynthesis in land plants. To address this issue, a gas exchange/chlorophyll fluorescence analysis was performed in three fern species spanning high phylogenetic range within Polypodiopsida (Osmunda regalis, Blechnum gibbum and Nephrolepis exaltata) to determine their maximum net photosynthesis (AN ), stomatal (gs ) and mesophyll (gm ) conductances to CO2 , and the maximum velocity of carboxylation (Vc,max ). The in vitro Rubisco specificity factor (SC /O ) was also determined. All three species had values for SC /O similar to those typical of seed plants, but values of AN , gs , gm and Vc,max were within the lowest range of those observed in seed plants. In addition, gs was unresponsive to light and CO2 , as already described in other fern species. On the contrary, gm varied with changes CO2 . A quantitative photosynthesis limitation analysis suggested that early land plants (ferns) presented not only stomatal limitations-which were less adjustable to the environment-but also restricted gm and Vc,max , resulting in limited maximum photosynthesis rates., (© 2013 Scandinavian Plant Physiology Society.)

Nothofagus nitida (Phil.) Krasser (Nothofagaceae) regenerates under the canopy in microsites protected from high light. Nonetheless, it is common to find older saplings in clear areas and adults as emergent trees of the Chilean evergreen forest. We hypothesized that this shade to sun transition in N. nitida is supported by an increase in photochemical and non-photochemical energy dissipation capacities of both photosystems in parallel with the increase in plant size and light availability. To dissect the relative contribution of light environment and plant developmental stage to these physiological responses, the photosynthetic performance of both photosystems was studied from the morpho-anatomical to the biochemical level in current-year leaves of N. nitida plants of different heights (ranging from 0.1 to 7 m) growing under contrasting light environments (integrated quantum flux (IQF) 5-40 mol m(-2). Tree height (TH) and light environment (IQF) independently increased the saturated electron transport rates of both photosystems, as well as leaf and palisade thickness, but non-photochemical energy flux, photoinhibition susceptibility, state transition capacity, and the contents of D1 and PsbS proteins were not affected by IQF and TH. Spongy mesophyll thickness and palisade cell diameter decreased with IQF and TH. A(max), light compensation and saturation points, Rubisco and nitrogen content (area basis) only increased with light environment (IQF), whereas dark respiration (R(d)) decreased slightly and relative chlorophyll content was higher in taller trees. Overall, the independent effects of more illuminated environment and tree height mainly increased the photochemical instead of the non-photochemical energy flux. Regardless of the photochemical increase with TH, carbon assimilation only significantly improved with higher IQF. Therefore it seems that mainly acclimation to the light environment supports the phenotypic transition of N. nitida from shade to sun.

Nothofagus nitida (Phil.) Krasser, an emergent tree of the Chilean evergreen forest, regenerates under the canopy. Nonetheless, it is common to find older saplings in clear areas. We hypothesized that this transition from shade to sun during the early developmental stages is made possible by an ontogenetic increase in the light acclimation capacity of photosynthesis. To test our hypothesis, we studied photosynthetic performance and photoprotection in N. nitida saplings at different developmental stages corresponding with three different height classes (short: 16.2 cm; medium-height: 48.0 cm; and tall: 73.7 cm) grown under contrasting light conditions (photosynthetic photon flux (PPF) of 20, 300 or 600 micromol m(-2) s(-1)) until newly expanded leaves had developed. Light-saturated CO(2) assimilation rate and light compensation and saturation points increased with increasing PPF. Medium-height and tall saplings acclimated to high light had higher electron transport rates and higher proportions of open Photosystem II reaction centers than shorter plants acclimated to high light. Short saplings showed higher thermal dissipation and contents of xanthophylls than taller saplings. Only medium-height and tall saplings acclimated to high light recovered after photoinhibition. State transitions were higher in short plants growing in low light, and decreased with plant height and growth irradiance. Thus, during development, N. nitida changes the balance of light energy utilization and photoprotective mechanisms, supporting a phenotypic transition from shade to sun during its early ontogeny.