http://digital.csic.es/bitstream/10261/60376/4/Mesophyll_diffusion_conductance.pdf

Mesophyll diffusion conductance to CO2: an unappreciated central player in photosynthesis

One of ecology's grand challenges is developing general rules to explain and predict highly complex systems. Understanding and predicting ecological processes from species' traits has been considered a 'Holy Grail' in ecology. Plant functional traits are increasingly being used to develop mechanistic models that can predict how ecological communities will respond to abiotic and biotic perturbations and how species will affect ecosystem function and services in a rapidly changing world; however, significant challenges remain. In this review, we highlight recent work and outstanding questions in three areas: (i) selecting relevant traits; (ii) describing intraspecific trait variation and incorporating this variation into models; and (iii) scaling trait data to community- and ecosystem-level processes. Over the past decade, there have been significant advances in the characterization of plant strategies based on traits and trait relationships, and the integration of traits into multivariate indices and models of community and ecosystem function. However, the utility of trait-based approaches in ecology will benefit from efforts that demonstrate how these traits and indices influence organismal, community, and ecosystem processes across vegetation types, which may be achieved through meta-analysis and enhancement of trait databases. Additionally, intraspecific trait variation and species interactions need to be incorporated into predictive models using tools such as Bayesian hierarchical modelling. Finally, existing models linking traits to community and ecosystem processes need to be empirically tested for their applicability to be realized.

/pdf/revisiting-the-holy-grail-using-plant-functional-traits-to-do0rqzowgt.pdf

Revisiting the Holy Grail: using plant functional traits to understand ecological processes

Plants are exposed to heterogeneity in the environment where new stress factors (i.e., climate change, land use change, and invasiveness) are introduced, and where inter- and intraspecies differences may reflect resource limitation and/or environmental stress factors. Phenotypic plasticity is considered one of the major means by which plants can cope with environmental factor variability. Nevertheless, the extent to which phenotypic plasticity may facilitate survival under environmental condition changes still remains largely unknown because results are sometimes controversial. Thus, it is important to identify plant functional traits in which plasticity may play a determinant role in plant response to global change as well as on the ecological consequences at an ecosystem level for the competition between wild and invasive species, considering that species with a greater adaptive plasticity may be more likely to survive in novel environmental conditions. In the near future, it will be important to increase long-term studies on natural populations in order to understand plant response to environmental factor fluctuations including climate change. There is the necessity to analyze variations at phenotypic and genetic levels for the same species and, in particular, for endemic and rare species because these could have drastic effects at an ecosystem level.

/pdf/plant-phenotypic-plasticity-in-response-to-environmental-3a7r62ja5v.pdf

Plant Phenotypic Plasticity in Response to Environmental Factors

The leaf economics spectrum (LES) represents a suite of intercorrelated leaf traits concerning construction costs per unit leaf area, nutrient concentrations, and rates of carbon fixation and tissue turnover. Although broad trade-offs among leaf structural and physiological traits have been demonstrated, we still do not have a comprehensive view of the fundamental constraints underlying the LES trade-offs. Here, we investigated physiological and structural mechanisms underpinning the LES by analysing a novel data compilation incorporating rarely considered traits such as the dry mass fraction in cell walls, nitrogen allocation, mesophyll CO2 diffusion and associated anatomical traits for hundreds of species covering major growth forms. The analysis demonstrates that cell wall constituents are major components of leaf dry mass (18-70%), especially in leaves with high leaf mass per unit area (LMA) and long lifespan. A greater fraction of leaf mass in cell walls is typically associated with a lower fraction of leaf nitrogen (N) invested in photosynthetic proteins; and lower within-leaf CO2 diffusion rates, as a result of thicker mesophyll cell walls. The costs associated with greater investments in cell walls underpin the LES: long leaf lifespans are achieved via higher LMA and in turn by higher cell wall mass fraction, but this inevitably reduces the efficiency of photosynthesis.

/pdf/physiological-and-structural-tradeoffs-underlying-the-leaf-3w8gzmawmx.pdf

Physiological and structural tradeoffs underlying the leaf economics spectrum.

A key objective for sustainable agriculture and forestry is to breed plants with both high carbon gain and water-use efficiency (WUE). At the level of leaf physiology, this implies increasing net photosynthesis (A N) relative to stomatal conductance (g s). Here, we review evidence for CO2 diffusional constraints on photosynthesis and WUE. Analyzing past observations for an extensive pool of crop and wild plant species that vary widely in mesophyll conductance to CO2 (g m), g s, and foliage A N, it was shown that both g s and g m limit A N, although the relative importance of each of the two conductances depends on species and conditions. Based on Fick's law of diffusion, intrinsic WUE (the ratio A N/g s) should correlate on the ratio g m/g s, and not g m itself. Such a correlation is indeed often observed in the data. However, since besides diffusion A N also depends on photosynthetic capacity (i.e., V c,max), this relationship is not always sustained. It was shown that only in a very few cases, genotype selection has resulted in simultaneous increases of both A N and WUE. In fact, such a response has never been observed in genetically modified plants specifically engineered for either reduced g s or enhanced g m. Although increasing g m alone would result in increasing photosynthesis, and potentially increasing WUE, in practice, higher WUE seems to be only achieved when there are no parallel changes in g s. We conclude that for simultaneous improvement of A N and WUE, genetic manipulation of g m should avoid parallel changes in g s, and we suggest that the appropriate trait for selection for enhanced WUE is increased g m/g s.

/pdf/diffusional-conductances-to-co2-as-a-target-for-increasing-3nxefz4chm.pdf

Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency.

In outdoor common gardens, high latitude populations of deciduous tree species often display higher assimilation rates (A) than low latitude populations, but they accomplish less height. To test whether trends in A reflect adaptation to growing season length or, alternatively, are garden growth artefacts, we examined variation in height increment and ecophysiological traits in a range-wide collection of Populus balsamifera L. populations from 21 provenances, during unconstrained growth in a greenhouse. Rooted cuttings, maintained without resource limitation under 21 h photoperiod for 90 d, displayed increasing height growth, A, leaf mass per area and leaf N per area with latitude whereas stomatal conductance (gs) showed no pattern. Water-use efficiency as indicated by both gas exchange and δ13C increased with latitude, whereas photosynthetic nitrogen-use efficiency decreased. Differences in δ13C were less than expected based on A/gs, suggesting coextensive variation in internal conductance (gm). Analysis of A–Ci curves on a subset of populations showed that high latitude genotypes had greater gm than low-latitude genotypes. We conclude that higher peak rates of height growth in high latitude genotypes of balsam poplar are supported by higher A, achieved partly through higher gm, to help compensate for a shorter growing season.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2009.02042.x

Enhanced assimilation rate and water use efficiency with latitude through increased photosynthetic capacity and internal conductance in balsam poplar (Populus balsamifera L.)

In plant ecophysiology, functional leaf traits are generally not assessed in relation to phenological phase of the canopy. Leaf traits measured in deciduous perennial species are known to vary between spring and summer seasons, but there is a knowledge gap relating to the late-summer phase marked by growth cessation and bud set occurring well before fall leaf senescence. The effects of phenology on canopy physiology were tested using a common garden of over 2,000 black cottonwood (Populus trichocarpa) individuals originating from a wide geographical range (44–60oN). Annual phenological events and 12 leaf-based functional trait measurements were collected spanning the entire summer season prior to, and following, bud set. Patterns of seasonal trait change emerged by synchronizing trees using their date of bud set. In particular, photosynthetic, mass, and N-based traits increased substantially following bud set. Most traits were significantly different between pre-bud set and post-bud set phase trees, with many traits showing at least 25 % alteration in mean value. Post-bud set, both the significance and direction of trait–trait relationships could be modified, with many relating directly to changes in leaf mass. In Populus, these dynamics in leaf traits throughout the summer season reflected a shift in whole plant physiology, but occurred long before the onset of leaf senescence. The marked shifts in measured trait values following bud set underscores the necessity to include phenology in trait-based ecological studies or large-scale phenotyping efforts, both at the local level and larger geographical scale.

Seasonality and phenology alter functional leaf traits

In outdoor common gardens, high latitude populations of deciduous tree species often display higher assimilation rates (A) than low latitude populations,but they accomplish less height.To test whether trends in A reflect adaptation to growing season length or, alternatively, are garden growth artefacts, we examined variation in height increment and ecophysiological traits in a range-wide collection of Populus balsamifera L. populations from 21 provenances, during unconstrained growth in a greenhouse. Rooted cut- tings, maintained without resource limitation under 21 h photoperiod for 90 d,displayed increasing height growth,A, leaf mass per area and leaf N per area with latitude whereas stomatal conductance (gs) showed no pattern. Water-use efficiency as indicated by both gas exchange and d 13 C increased with latitude, whereas photosynthetic nitrogen- use efficiency decreased. Differences in d 13 C were less than expected based on A/gs, suggesting coextensive variation in internal conductance (gm). Analysis of A-Ci curves on a subset of populations showed that high latitude genotypes had greater gm than low-latitude genotypes. We conclude that higher peak rates of height growth in high latitude genotypes of balsam poplar are supported by higher A, achieved partly through higher gm,to help compensate for a shorter growing season.

Eric C. Drewes

Papers

Enhanced assimilation rate and water use efficiency with latitude through increased photosynthetic capacity and internal conductance in balsam poplar (Populus balsamifera L.)

Seasonality and phenology alter functional leaf traits

Enhanced assimilation rate and water use efficiency with latitude through increased photosynthetic capacity and internal conductance in balsam poplar