Protocol

Author

This article is modified from Perez-Harguindeguy et al (2013). The “New handbook for standardised measurement of plant functional traits worldwide” is a product of and is hosted by Nucleo Diversus (with additional Spanish translation). For more on this trait and on its context as part of the entire trait handbook visit its primary site Nucleo DiverSus at http://www.nucleodiversus.org/?lang=en

Contributing Author

Louis Santiago

Summary

The light-saturated photosynthetic rate (A_max) under typical field conditions, usually expressed in μmol m^-2 s^-1 or nmol g^-1 s^-1, or both, is a valuable metric as a measure (or at least as an index) of metabolic capacity and a factor determining average realised photosynthetic rate (for upper-canopy foliage). A_max scales with other structural, chemical and longevity aspects of the leaf economic spectrum and, along with those other variables, enables scaling to canopy processes of whole ecosystems. Simultaneous measures of leaf water-vapour conductance are typically made in concert with the photosynthetic measurements.

What, when and how to measure

Sample young, fully expanded leaves (see SLA). These should be from sunlit parts of the canopy, unless specifically focusing on the shaded taxa of understorey. Measure leaves only if they have been in sufficiently high light just before measurement (e.g. direct sun for 5-10 min) to minimise concerns about leaf induction status or stomatal closure as a result of shading (see SLA for discussion).

Because realised photosynthesis is less than maximal because of a host of factors, including low or high temperatures, limited soil moisture or air humidity, negative leaf water potential, and source-sink inhibition, among others, care must be taken in choosing the time of year, time of day, and general conditions under which measurements can be made. Some knowledge of gas-exchange responses of the taxa under study will be essential. Do not make measurements during or just following (days to weeks) periods of severe water deficit, or unusual temperatures. Do make measurements on days when soil moisture, plant water status, air humidity, irradiance and temperatures are near optimal for the taxa in question. Measurements in most ecosystems should be made at mid- to late morning (e.g. from 0800 hours to 1100 hours local time) under non-limiting vapour-pressure deficits or temperatures. This minimises the risk of sampling during midday and afternoon declines in gas-exchange rate as a result of stomatal closure, source-sink inhibition or other causes. If a given morning, or mornings in general, are cold relative to photosynthetic temperature optima, measurement can be made later in the day. Because most published measurements have been made under ambient CO₂ concentrations, that would be recommended. If rates can also be measured under saturating CO₂ concentrations, that is also useful.

Any reliable leaf gas-exchange system can be used, and conditions in the chamber can be either set at levels considered optimal or left to track the in situ conditions (which need to be near optimal). If possible, measure intact foliage or else leaves on branches cut and then re-cut underwater. In the latter case, check whether given individuals fail to stay hydrated. Conduct some test comparisons of gas exchange on intact and -re-cut’ branches, to ensure the technique works for your taxa and system. Measurements can also be made on detached foliage; however, this requires even greater attention. Leaves should be measured within seconds, to a few minutes at most, after detachment, and tests of intact v. detached foliage should be made for a subsample, to ensure similar rates are observed.

If possible, the leaf material inside the chamber should be collected (see SLA), measured for LDMC and SLA, and stored for any subsequent chemical analyses.

Further information

For further information and protocols on the analysis of instantaneous and maximum photosynthetic rate data collected, see Instantaneous gas exchange data, in the Statistics section.

Literature references

References on theory, significance and large datasets:

Reich PB, Walters MB, Ellsworth DS (1992) Leaf life-span in relation to leaf, plant and stand characteristics among diverse ecosystems. Ecological Monographs 62, 365-392 doi:102307/2937116

Reich PB, Walters MB, Ellsworth DS (1997) From tropics to tundra: global convergence in plant functioning. Proceedings of the National Academy of Sciences, USA 94, 13730-13734 doi:101073/pnas942513730

More on methods:

Ellsworth DS, Reich PB (1992) Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes. Tree Physiology 10, 1-20 doi:101093/treephys/1011

Reich PB, Walters MB, Ellsworth DS (1991) Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees. Plant, Cell & Environment 14, 251-259 doi:101111/j1365-30401991tb01499x

Santiago LS, Wright SJ (2007) Leaf functional traits of tropical forest plants in relation to growth form. Functional Ecology 21:19-27

Wong S-C, Cowan IR, Farquhar GD (1985b) Leaf conductance in relation to rate of CO₂ assimilation. II. Effects of short-term exposures to different photon flux densities. Plant Physiology 78, 826-829 doi:101104/pp784826

Wong S-C, Cowan IR, Farquhar GD (1985c) Leaf conductance in relation to rate of CO₂assimilation. III. Influences of water stress and photoinhibition. Plant Physiology 78, 830-834 doi:101104/pp784830