Summary
Large variations are found in leaf morphology and physiology across species in nature, reflecting diversity in carbon fixation and growth strategies. These variations in leaf traits are not random; rather, they are tightly coordinated with each other. Leaf traits can be expressed per leaf dry mass or per leaf area. A leaf-mass basis reflects leaf economics, i.e., revenues and expenditures per unit investment of biomass, while a leaf-area basis reflects fluxes in relation to surfaces. Leaf N and P concentrations, and photosynthetic and respiration rates – all considered on a mass basis, are negatively correlated with leaf mass per area (LMA) whilst leaf lifespan is positively correlated with LMA. These correlations are summarized into a single major axis called the “leaf economics spectrum” that runs from “quick-return” to “slow-return” species. On the other hand, correlations among area-based traits are less consistent and less understood in relation to leaf economy. LMA was positively correlated with leaf N content but mostly independent from photosynthetic rates per unit leaf area. Given that N is an essential element in photosynthetic proteins and thus photosynthesis, clarifying the mechanisms why the efficiency of photosynthesis (photosynthesis per unit N) decreases with LMA is a major concern in understanding the correlations among area-based traits in relation to leaf economy. Currently available data suggest that greater amounts of cell wall are required for long-lived leaves, which reduces the efficiency of photosynthesis by lowering (1) the fraction of leaf N invested in photosynthetic proteins and (2) CO2 diffusion rates through thicker and denser mesophyll cell walls. These physiological and structural constraints are a fundamental mechanism underpinning the general correlations among leaf economic traits.
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Abbreviations
- A:
-
assimilation rate
- Aarea :
-
net assimilation rate per unit leaf area
- Amass :
-
net assimilation rate per unit leaf dry mass
- C:
-
carbon
- Ca :
-
ambient CO2 concentration
- Cc :
-
chloroplast CO2 concentration
- Ci :
-
intercellular CO2 concentration
- CWarea :
-
cell wall mass per unit leaf area
- CWmass :
-
cell wall mass per unit leaf mass
- Dec:
-
deciduous
- DM:
-
dry mass
- Eve:
-
evergreen
- gm :
-
mesophyll conductance for CO2
- gs :
-
stomatal conductance for CO2
- LES:
-
leaf economics spectrum
- LL:
-
leaf lifespan
- LMA:
-
leaf mass per area
- N:
-
nitrogen
- Narea :
-
leaf nitrogen (N) content per unit leaf area
- Nmass :
-
leaf N concentration
- Np :
-
photosynthetic N content per unit leaf area
- NRub :
-
rubisco N content per unit leaf area
- P:
-
phosphorus
- PNUE:
-
photosynthetic N use efficiency measured at saturating light intensity (=Aarea/Narea)
- QR:
-
quantile range
- R:
-
respiration rate in the dark
- Rarea :
-
dark respiration rate per unit leaf area
- Rmass :
-
dark respiration rate per unit leaf dry mass
- Rubisco:
-
ribulose bisphosphate carboxylase oxygenase
- Sc :
-
surface area of chloroplasts exposed to intercellular airspace per unit leaf area
- Sm :
-
surface area of mesophyll exposed to intercellular airspace per unit leaf area
- SMA:
-
standardized major axis
- TCW :
-
mesophyll cell wall thickness
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This study is partly supported by grants from JSPS KAKENHI #26711025 (YO), and from the Australian Research Council (IW).
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Onoda, Y., Wright, I.J. (2018). The Leaf Economics Spectrum and its Underlying Physiological and Anatomical Principles. In: Adams III, W., Terashima, I. (eds) The Leaf: A Platform for Performing Photosynthesis. Advances in Photosynthesis and Respiration, vol 44. Springer, Cham. https://doi.org/10.1007/978-3-319-93594-2_16
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