Specific Leaf Area
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Specific Leaf Area

Specific leaf area (SLA) is the ratio of leaf area to leaf dry mass.[1][2][3] The inverse of SLA is Leaf Mass per Area (LMA).

## Rationale

Specific leaf area is a ratio indicating how much leaf area a plant builds with a given amount of leaf biomass:

${\displaystyle SLA\ =\ {\frac {A}{M_{L}}}}$

where A is the area of a given leaf or all leaves of a plant, and ML is the dry mass of those leaves. Typical units are m2.kg-1 or mm2.mg-1.

Leaf mass per area (LMA) is its inverse and can mathematically be decomposed in two component variables, leaf thickness (LTh) and leaf density (LD):[4]

${\displaystyle LMA\ =\ {\frac {M_{L}}{A}}\ =LTh.LD}$

Typical units are g.m-2 for LMA, µm for LTh and g.ml-1 for LD.

Both SLA and LMA are frequently used in plant ecology and biology. SLA is one of the components in plant growth analysis, and mathematically scales positively and linearly with the relative growth rate of a plant. LMA mathematically scales positively with the investments plants make per unit leaf area (amount of protein and cell wall; cell number per area) and with leaf longevity. Since linear, positive relationships are more easily analysed than inverse negative relationships, researchers often use either variable, depending on the type of questions asked.

## Normal Ranges

Normal ranges of SLA and LMA are species-dependent and influenced by growth environment. Table 1 gives normal ranges (~10th and ~90th percentiles) for species growing in the field, for well-illuminated leaves. Aquatic plants generally have very low LMA values, with particularly low numbers reported for species such as Myriophyllum farwelli (2.8 g.m-2)[5] and Potamogeton perfoliatus (3.9 g. m-2).[6] Evergreen shrubs and Gymnosperm trees as well as succulents have particularly high LMA values, with highest values reported for Aloe saponaria (2010 g.m-2) and Agave deserti (2900 g.m-2).[7]

Table 1. Normal ranges of SLA and LMA for plant species growing in the field.[8]
SLA LMA
Normal Range low high low high
Herbaceous

species

Aquatic plants 15 210 4 65
Ferns 10 45 20 95
Forbs 8 35 25 130
Grasses 4 30 35 225
Succulents 2 12 85 510
Woody

species

Deciduous shrubs 9 27 35 110
Deciduous woody species 8 25 40 120
Evergreen shrubs 2 15 65 380
Evergreen Angiosperm trees 8 23 40 120
Evergreen Gymnosperm trees 2 11 90 470

## Application

Specific leaf area can be used to estimate the reproductive strategy of a particular plant based upon light and moisture (humidity) levels, among other factors.[9] Specific leaf area is one of the most widely accepted key leaf characteristics used during the study of leaf traits.[10][11][12][13]

## Changes in response to drought

Drought and water stress have varying effects on specific leaf area. In a variety of species, drought decreases specific leaf area.[14][15][16] For example, under drought conditions, leaves were, on average, smaller than leaves on control plants.[17] This is a logical observation, as a relative decrease in surface area would mean that there would be fewer ways for water to be lost. Species with typically low specific leaf area values are geared for the conservation of acquired resources, due to their large dry matter content, high concentrations of cell walls and secondary metabolites, and high leaf and root longevity.[18]

In some other species, such as Poplar trees, specific leaf area will decrease overall, but there will be an increase in specific leaf area until the leaf has reached its final size. After the final size has been reached, the specific leaf area will then begin decreasing.[19]

Other research has shown increasing specific leaf area values in plants under water limitation. An example of increasing specific leaf area values as a result of drought stress is the birch tree species.[20] Birch tree specific leaf area values significantly increased after two dry seasons, though the authors did note that, in typical cases, lowered specific leaf area values are seen as an adaptation to drought stress.

## References

1. ^ Vile, D.; Garnier, E; Shipley, B; Laurent, G; Navas, ML; Roumet, C; Lavorel, S; Díaz, S; et al. (2005). "Specific Leaf Area and Dry Matter Content Estimate Thickness in Laminar Leaves". Annals of Botany. 96 (6): 1129-1136. doi:10.1093/aob/mci264. PMC 4247101. PMID 16159941. Retrieved .
2. ^ "Specific leaf area and leaf dry matter content of plants growing in sand dunes". Ejournal.sinica.edu.tw. Retrieved . Cite journal requires |journal= (help)
3. ^ "Varietal differences in specific leaf area: a common physiological determinant of tillering ability and early growth vigor ? - Publications des agents du Cirad". Publications.cirad.fr. Retrieved . Cite journal requires |journal= (help)
4. ^ Witkowski, E. T. F.; Lamont, Byron B. (1991-12-01). "Leaf specific mass confounds leaf density and thickness". Oecologia. 88 (4): 486-493. doi:10.1007/BF00317710. ISSN 1432-1939. PMID 28312617.
5. ^ Gerber, D. Timothy; Les, Donald H. (1994). "Comparison of leaf morphology among submersed species of Myriophyllum (Haloragaceae) from different habitats and geographical distributions". American Journal of Botany. 81 (8): 973-979. doi:10.1002/j.1537-2197.1994.tb15584.x. ISSN 1537-2197.
6. ^ Neundorfer, JV; Kemp, WM (1993). "Nitrogen versus phosphorus enrichment of brackish waters: responses of the submersed plant Potamogeton perfoliatus and its associated algal community". Marine Ecology Progress Series. 94: 71-82. doi:10.3354/meps094071. ISSN 0171-8630.
7. ^ Turner, Neil C, editor literario. Kramer, Paul Jackson, editor literario. (1980). Adaptation of plants to water and high temperature stress. John Wiley & Sons. ISBN 0471053724. OCLC 932308683.CS1 maint: multiple names: authors list (link)
8. ^ Poorter, Hendrik; Niinemets, Ülo; Poorter, Lourens; Wright, Ian J.; Villar, Rafael (2009). "Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis". New Phytologist. 182 (3): 565-588. doi:10.1111/j.1469-8137.2009.02830.x. ISSN 1469-8137. PMID 19434804.
9. ^ Milla, R.; Reich P. B. (4 Apr 2008). "Environmental and developmental controls on specific leaf area are little modified by leaf allometry". Functional Ecology. 22 (4): 565-576. doi:10.1111/j.1365-2435.2008.01406.x. Archived from the original on 5 January 2013. Retrieved 2009.
10. ^ Freschet G.T., Dias A.T.C., Ackerly D.D., Aerts R., van Bodegom P.M., Cornwell W.K., Dong M., Kurokawa H., Liu G., Onipchenko V.G., Ordoñez J.C., Peltzer D.A., Richardson S.J., Shidakov I.I., Soudzilovskaia N.A., Tao J. & Cornelissen J.H.C. (2011). Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages. Global Ecology and Biogeography, no-no.
11. ^ Hoffmann, W.A.; Franco, A.C.; Moreira, M.Z.; Haridasan, M. (2005). "Specific leaf area explains differences in leaf traits between congeneric savanna and forest trees". Functional Ecology. 19 (6): 932-940. doi:10.1111/j.1365-2435.2005.01045.x.
12. ^ Kraft, N.J.B.; Valencia, R.; Ackerly, D.D. (2008). "Functional traits and niche-based tree community assembly in an amazonian forest". Science. 322 (5901): 580-582. doi:10.1126/science.1160662. PMID 18948539.
13. ^ Wright, I.J.; Reich, P.B.; Westoby, M.; Ackerly, D.D.; Baruch, Z.; Bongers, F.; Cavender-Bares, J.; Chapin, T.; Cornelissen, J.H.C.; Diemer, M.; Flexas, J.; Garnier, E.; Groom, P.K.; Gulias, J.; Hikosaka, K.; Lamont, B.B.; Lee, T.; Lee, W.; Lusk, C.; Midgley, J.J.; Navas, M.-L.; Niinemets, Ü.; Oleksyn, J.; Osada, N.; Poorter, H.; Poot, P.; Prior, L.; Pyankov, V.I.; Roumet, C.; Thomas, S.C.; Tjoelker, M.G.; Veneklass, E.J.; Villar, R. (2004). "The worldwide leaf economics spectrum". Nature. 428 (6985): 821-827. doi:10.1038/nature02403. hdl:11299/176900. PMID 15103368.
14. ^ Casper, B. B.; Forseth, I. N.; Kempenich, H.; Seltzer, S.; Xavier, K. (2001). "Drought prolongs leaf life span in the herbaceous desert perennial Cryptantha flava". Functional Ecology. 15 (6): 740-747. doi:10.1046/j.0269-8463.2001.00583.x.
15. ^ Marron, N.; Dreyer, E.; Boudouresque, E.; Delay, D.; Petit, J.-M.; Delmotte, F. M.; Brignolas, F. (2003). "Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus canadensis (Moench) clones,"Dorskamp"and "Luisa_Avanzo"". Tree Physiology. 23 (18): 1225-1235. doi:10.1093/treephys/23.18.1225. PMID 14652222.
16. ^ Laureano, R. G.; Lazo, Y. O.; Linares, J. C.; Luque, A.; Martínez, F.; Seco, J. I.; Merino, J. (2008). "The cost of stress resistance: construction and maintenance costs of leaves and roots in two populations of Quercus ilex". Tree Physiology. 28 (11): 1721-1728. doi:10.1093/treephys/28.11.1721. PMID 18765377.
17. ^ Casper, B. B.; Forseth, I. N.; Kempenich, H.; Seltzer, S.; Xavier, K. (2001). "Drought prolongs leaf life span in the herbaceous desert perennial Cryptantha flava". Functional Ecology. 15 (6): 740-747. doi:10.1046/j.0269-8463.2001.00583.x.
18. ^ Marron, N.; Dreyer, E.; Boudouresque, E.; Delay, D.; Petit, J.-M.; Delmotte, F. M.; Brignolas, F. (2003). "Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus canadensis (Moench) clones,"Dorskamp"and "Luisa_Avanzo"". Tree Physiology. 23 (18): 1225-1235. doi:10.1093/treephys/23.18.1225. PMID 14652222.
19. ^ Marron, N.; Dreyer, E.; Boudouresque, E.; Delay, D.; Petit, J.-M.; Delmotte, F. M.; Brignolas, F. (2003). "Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus canadensis (Moench) clones,"Dorskamp"and "Luisa_Avanzo"". Tree Physiology. 23 (18): 1225-1235. doi:10.1093/treephys/23.18.1225. PMID 14652222.
20. ^ Aspelmeier, S.; Leuschner, C. (2006). "Genotypic variation in drought response of silver birch (Betula pendula Roth): leaf and root morphology and carbon partitioning". Trees. 20: 42-52. doi:10.1007/s00468-005-0011-9.