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Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE

Different responses in leaf pigments and leaf mass per area to altitude between evergreen and deciduous woody species

Yan Li A , Dongmei Yang B D , Shuang Xiang C and Guoyong Li C
+ Author Affiliations
- Author Affiliations

A Department of Biology, Nanjing University, 22 Hankou Road, Nanjing 210093, China.

B College of Chemistry and Life Sciences, Zhejiang Normal University, 688 Yingbin Road, Jinhua 321004, China.

C Center for Ecological Research, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China.

D Corresponding author. Email: yangdm@zjnu.cn

Australian Journal of Botany 61(6) 424-435 https://doi.org/10.1071/BT13022
Submitted: 30 January 2013  Accepted: 12 July 2013   Published: 21 August 2013

Abstract

Leaf chlorophyll content is positively associated with photosynthetic capacity and nutrient status, but its functional ecology has seldom been examined thus far. In the present study, we measured leaf chlorophyll and carotenoid concentrations, determined chlorophyll a : chlorophyll b (Chl a : Chl b) and carotenoids : chlorophyll ratios and measured leaf mass per area (LMA) for 63 woody dicot species, including 24 evergreen species and 39 deciduous species, at two altitudes (1800–2400 and 2400–2800 m a.s.l.) of Gongga Mountain, south-west China. The aim of the present study was to determine whether evergreen and deciduous species differ in terms of leaf pigment concentrations and LMA in response to environmental differences associated with changes in elevation. In both life forms, the altitude effect was not significant for chlorophyll and carotenoid concentrations. However, the Chl a : Chl b ratio was significantly higher in evergreen species, whereas LMA was significantly higher in deciduous species, at the high versus low altitude. These observations suggest that evergreen and deciduous species may have different strategies to protect leaf pigments. Mass-based leaf pigment concentrations were lower in evergreen compared with deciduous species, especially at high altitude. LMA was higher in evergreen than deciduous species at both altitudes. Pigment concentrations were negatively correlated with LMA in both life forms at both altitudes. The slope of LMA vs mass-based leaf pigment concentrations was significantly more negative for deciduous than evergreen species, and at low versus high altitude for deciduous species. The findings suggest that deciduous species may invest less photosynthate in leaf pigments but more in inactive components in stressful environments than do evergreens. Thus, the same magnitude of variation in LMA may have different consequences on leaf carbon balance between evergreen and deciduous species, which helps explain why evergreen species are often more likely to occupy more stressful environments than deciduous species.

Additional keywords: chlorophyll, carotenoids, environmental gradient, life form, life history strategy.


References

Atkin OK, Loveys BR, Atkinson LJ, Pons TL (2006) Phenotypic plasticity and growth temperature: understanding interspecific variability. Journal of Experimental Botany 57, 267–281.
Phenotypic plasticity and growth temperature: understanding interspecific variability.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28XisFKnsg%3D%3D&md5=e29d100459080dcf09b40c45f843e084CAS | 16371402PubMed |

Caldwell MM, Robberecht R, Flint SD (1983) Internal filters: prospects of UV-acclimation in higher plants. Physiologia Plantarum 58, 445–450.
Internal filters: prospects of UV-acclimation in higher plants.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL3sXks1KktL4%3D&md5=4dab98ce7d09f8961b22a2bf0ddde4b2CAS | 1:CAS:528:DyaL3sXks1KktL4%3D&md5=4dab98ce7d09f8961b22a2bf0ddde4b2CAS |

Castro-Díez P, Puyravaud J, Cornelissen J (2000) Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types. Oecologia 124, 476–486.
Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types.Crossref | GoogleScholarGoogle Scholar |

Chen T, Zhao Z, Zhang Y, Qiang W, Feng H, An L, Li Z (2012) Physiological variations in chloroplasts of Rhodiola coccinea along an altitudinal gradient in Tianshan Mountain. Acta Physiologiae Plantarum 34, 1007–1015.
Physiological variations in chloroplasts of Rhodiola coccinea along an altitudinal gradient in Tianshan Mountain.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XhtVKntrvM&md5=14e749ecbe0527e3a2eca4b97e285720CAS |

Cheng G, Luo J (2002) Successional features and dynamic simulation of sub-alpine forest in the Gongga Mountain China. Acta Ecologica Sinica 22, 1049–1056. [in Chinese]

Cordell S, Goldstein G, Meinzer FC, Handley LL (1999) Allocation of nitrogen and carbon in leaves of Metrosideros polymorpha regulates carboxylation capacity and 13C along an altitudinal gradient. Functional Ecology 13, 811–818.
Allocation of nitrogen and carbon in leaves of Metrosideros polymorpha regulates carboxylation capacity and 13C along an altitudinal gradient.Crossref | GoogleScholarGoogle Scholar |

De Lillis M, Matteucci G, Valentini R (2004) Carbon assimilation, nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient. Photosynthetica 42, 597–605.
Carbon assimilation, nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXhsVOmtLg%3D&md5=3cc39f65925d7b8e7b5583fb576b51ffCAS |

Demmig-Adams B, Adams W (1996) The role of xanthophyll cycle carotenoids in the protection of photosynthesis. Trends in Plant Science 1, 21–26.
The role of xanthophyll cycle carotenoids in the protection of photosynthesis.Crossref | GoogleScholarGoogle Scholar |

Emerson R (1929) Chlorophyll content and rate of photosynthesis. Proceedings of the National Academy of Sciences of the United States of America 15, 281–284.
Chlorophyll content and rate of photosynthesis.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaB1MXjtFClsQ%3D%3D&md5=9e7779e33fe52a1816ede521b918418dCAS | 16577182PubMed |

Falster DS, Warton DI, Wright IJ (2006). ‘User’s guide to SMATR: Standardised Major Axis Tests & Routines, Version 2.0.’ Available at: http://www.bio.mq.edu.au/ecology/SMATR/ [Verified 11 March 2006].

Filella I, Peñuelas J (1999) Altitudinal differences in UV absorbance, UV reflectance and related morphological traits of Quercus ilex and Rhododendron ferrugineum in the Mediterranean region. Plant Ecology 145, 157–165.
Altitudinal differences in UV absorbance, UV reflectance and related morphological traits of Quercus ilex and Rhododendron ferrugineum in the Mediterranean region.Crossref | GoogleScholarGoogle Scholar |

Friend AD, Woodward FI, Switsur VR (1989) Field measurements of photosynthesis, stomatal conductance, leaf nitrogen and δ13C along altitudinal gradients in Scotland. Functional Ecology 3, 117–122.
Field measurements of photosynthesis, stomatal conductance, leaf nitrogen and δ13C along altitudinal gradients in Scotland.Crossref | GoogleScholarGoogle Scholar |

Gilmore A (1997) Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves. Physiologia Plantarum 99, 197–209.
Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2sXjtFWrt70%3D&md5=e1dedb43588677f9f084061440466317CAS | 1:CAS:528:DyaK2sXjtFWrt70%3D&md5=e1dedb43588677f9f084061440466317CAS |

Godnev T, Hodasevic E (1965) Biosynthesis of pigments in some evergreen plants at temperatures below 0°C. Doklady Akademii Nauk SSSR 160, 1206–1208. [in Russian with an English abstract]

González J, Gallardo M, Boero C, Cruz ML, Prado F (2007) Altitudinal and seasonal variation of protective and photosynthetic pigments in leaves of the world’s highest elevation trees Polylepis tarapacana (Rosaceae). Acta Oecologica 32, 36–41.
Altitudinal and seasonal variation of protective and photosynthetic pigments in leaves of the world’s highest elevation trees Polylepis tarapacana (Rosaceae).Crossref | GoogleScholarGoogle Scholar |

Hacke UG, Sperry JS, Pockman WT, Davis SD, McCulloh KA (2001) Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure. Oecologia 126, 457–461.
Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure.Crossref | GoogleScholarGoogle Scholar |

Haldimann P (1996) Effects of changes in growth temperature on photosynthesis and carotenoid composition in Zea mays leaves. Physiologia Plantarum 97, 554–562.
Effects of changes in growth temperature on photosynthesis and carotenoid composition in Zea mays leaves.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK28Xktl2gtrg%3D&md5=ac2e863ada6f448e3ca26d5ff932bdb8CAS |

Hallik L, Kull O, Niinemets Ü, Aan A (2009) Contrasting correlation networks between leaf structure, nitrogen and chlorophyll in herbaceous and woody canopies. Basic and Applied Ecology 10, 309–318.
Contrasting correlation networks between leaf structure, nitrogen and chlorophyll in herbaceous and woody canopies.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXovFSgu7o%3D&md5=75ba1305b99ed5bd603856e089521e76CAS |

Harvey PJ, Pagel MD (1991). ‘The comparative method in evolutionary biology.’ (Oxford University Press: Oxford)

Hikosaka K (2004) Interspecific difference in the photosynthesis–nitrogen relationship: patterns, physiological causes, and ecological importance. Journal of Plant Research 117, 481–494.
Interspecific difference in the photosynthesis–nitrogen relationship: patterns, physiological causes, and ecological importance.Crossref | GoogleScholarGoogle Scholar | 15583974PubMed |

Huner N, Öquist G, Hurry V, Krol M, Falk S, Griffith M (1993) Photosynthesis, photoinhibition and low temperature acclimation in cold tolerant plants. Photosynthesis Research 37, 19–39.
Photosynthesis, photoinhibition and low temperature acclimation in cold tolerant plants.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3sXmsVGnurY%3D&md5=81beff75a075bace6c3384553cc68695CAS |

Kan K, Thornber J (1976) The light-harvesting chlorophyll a/b–protein complex of Chlamydomonas reinhardii. Plant Physiology 57, 47–52.
The light-harvesting chlorophyll a/b–protein complex of Chlamydomonas reinhardii.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE28Xos12mug%3D%3D&md5=7896fa6d001f29a77295a8beed0328ebCAS | 16659422PubMed |

Kao W, Chang K (2001) Altitudinal trends in photosynthetic rate and leaf characteristics of Miscanthus populations from central Taiwan. Australian Journal of Botany 49, 509–514.
Altitudinal trends in photosynthetic rate and leaf characteristics of Miscanthus populations from central Taiwan.Crossref | GoogleScholarGoogle Scholar |

Karabourniotis G, Papadopoulos K, Papamarkou M, Manetas Y (1992) Ultraviolet-B radiation absorbing capacity of leaf hairs. Physiologia Plantarum 86, 414–418.
Ultraviolet-B radiation absorbing capacity of leaf hairs.Crossref | GoogleScholarGoogle Scholar |

Körner C (2007) The use of ‘altitude’ in ecological research. Trends in Ecology & Evolution 22, 569–574.
The use of ‘altitude’ in ecological research.Crossref | GoogleScholarGoogle Scholar |

Körner C, Diemer M (1987) In situ photosynthetic responses to light, temperature and carbon dioxide in herbaceous plants from low and high altitude. Functional Ecology 1, 179–194.
In situ photosynthetic responses to light, temperature and carbon dioxide in herbaceous plants from low and high altitude.Crossref | GoogleScholarGoogle Scholar |

Körner C, Diemer M (1994) Evidence that plants from high altitudes retain their greater photosynthetic efficiency under elevated CO2. Functional Ecology 8, 58–68.
Evidence that plants from high altitudes retain their greater photosynthetic efficiency under elevated CO2.Crossref | GoogleScholarGoogle Scholar |

Lambers H, Poorter H (1992) Inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences. Advances in Ecological Research 23, 187–261.
Inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3sXksVGiu7w%3D&md5=07a20255b50b7a30d2fe780709b7e2c2CAS | 1:CAS:528:DyaK3sXksVGiu7w%3D&md5=07a20255b50b7a30d2fe780709b7e2c2CAS |

Lefsrud M, Kopsell A (2006) Biomass production and pigment accumulation in kale grown under different radiation cycles in a controlled environment. HortScience 41, 1412–1415.

Li GY, Yang DM, Sun SC (2008) Allometric relationships between lamina area, lamina mass and petiole mass of 93 temperate woody species vary with leaf habit, leaf form and altitude. Functional Ecology 22, 557–564.
Allometric relationships between lamina area, lamina mass and petiole mass of 93 temperate woody species vary with leaf habit, leaf form and altitude.Crossref | GoogleScholarGoogle Scholar |

Linder S (1971) Photosynthetic action spectra of Scots pine needles of different ages from seedlings grown under different nursery conditions. Physiologia Plantarum 25, 58–63.
Photosynthetic action spectra of Scots pine needles of different ages from seedlings grown under different nursery conditions.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE3MXltVKhtb8%3D&md5=f6ac8dbfba9b83d6a17815ec3162caa9CAS |

Liu Z (1985). ‘Vegetation of Gongga Mountain.’ (Sichuan Science and Technology Press: Chengdu) [in Chinese]

Lloyd J, Bloomfield K, Domingues TF, Farquhar GD (2013) Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: of ecophysiologcical relevance or just a case of mathematical imperatives and statistical quicksand? New Phytologist 199, 311–321.
Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: of ecophysiologcical relevance or just a case of mathematical imperatives and statistical quicksand?Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXpvFars7s%3D&md5=65ce0f440b54f0da2113c8f197d36afdCAS | 1:CAS:528:DC%2BC3sXpvFars7s%3D&md5=65ce0f440b54f0da2113c8f197d36afdCAS | 23621613PubMed |

Martins EP (2004). ‘COMPARE, version 4.6b. Computer programs for the statistical analysis of comparative data.’ Available at: http://compare.bio.indiana.edu/ [Verified 12 March 2011].

Matsubara S, Krause GH, Aranda J, Virgo A, Beisel KG, Jahns P, Winter K (2009) Sun–shade patterns of leaf carotenoid composition in 86 species of neotropical forest plants. Functional Plant Biology 36, 20–36.
Sun–shade patterns of leaf carotenoid composition in 86 species of neotropical forest plants.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXhs1GgsA%3D%3D&md5=2232c5419f96e64300ef008f50d820dcCAS |

Middleton E, Teramura A (1993) The role of flavonol glycosides and carotenoids in protecting soybean from ultraviolet-B damage. Plant Physiology 103, 741–752.

Munné-Bosch S, Alegre L (2000) Changes in carotenoids, tocopherols and diterpenes during drought and recovery, and the biological significance of chlorophyll loss in Rosmarinus officinalis plants. Planta 210, 925–931.
Changes in carotenoids, tocopherols and diterpenes during drought and recovery, and the biological significance of chlorophyll loss in Rosmarinus officinalis plants.Crossref | GoogleScholarGoogle Scholar | 10872224PubMed |

Neilson R, Ludlow M, Jarvis P (1972) Photosynthesis in sitka spruce (Picea sitchensis (Bong.) Carr.). II. Response to temperature. Journal of Applied Ecology 9, 721–745.
Photosynthesis in sitka spruce (Picea sitchensis (Bong.) Carr.). II. Response to temperature.Crossref | GoogleScholarGoogle Scholar |

Osnas JLD, Lichstein JW, Reich PB, Pacala SW (2013) Global leaf tait relationships: mass, area, and the leaf economics spectrum. Science 340, 741–744.
Global leaf tait relationships: mass, area, and the leaf economics spectrum.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXntVGjsLg%3D&md5=1b8332c544893b02f5fb6eb703617e52CAS |

Owens TG (1996). Processing of excitation energy by antenna pigments. In ‘Photosynthesis and the environment’. (Ed. NR Baker) pp. 1–21. (Kluwer Academic Publisher: NewYork)

Poorter H, Pepin S, Rijkers T, Jong Y, Evans J, Körner C (2006) Construction costs, chemical composition and payback time of high- and low-irradiance leaves. Journal of Experimental Botany 57, 355–371.
Construction costs, chemical composition and payback time of high- and low-irradiance leaves.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28XisFKgsQ%3D%3D&md5=5328b23341fc0ee7dbed176c1cbe566eCAS | 16303828PubMed |

Poorter H, Niinemets Ü, Poorter L, Wright IJ, Villar R (2009) Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytologist 182, 565–588.
Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis.Crossref | GoogleScholarGoogle Scholar | 19434804PubMed |

Reich PB, Walters MB, Ellsworth DS (1997) From tropics to tundra: global convergence in plant functioning. Proceedings of the National Academy of Sciences of the United States of America 94, 13 730–13 734.
From tropics to tundra: global convergence in plant functioning.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2sXotValtb8%3D&md5=ee7a5edd16fa9ff019ef52afe2ee29e3CAS |

Reich PB, Oleksyn J, Wright IJ (2009) Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species. Oecologia 160, 207–212.
Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species.Crossref | GoogleScholarGoogle Scholar | 19212782PubMed |

Robinson SA, Turnbull JD, Lovelock CE (2005) Impact of changes in natural UV radiation on pigment composition, physiological and morphological characteristics of the Antarctic moss, Grimmia antarctici. Global Change Biology 11, 476–489.
Impact of changes in natural UV radiation on pigment composition, physiological and morphological characteristics of the Antarctic moss, Grimmia antarctici.Crossref | GoogleScholarGoogle Scholar |

Rosevear M, Young A, Johnson G (2001) Growth conditions are more important than species origin in determining leaf pigment content of British plant species. Functional Ecology 15, 474–480.
Growth conditions are more important than species origin in determining leaf pigment content of British plant species.Crossref | GoogleScholarGoogle Scholar |

Rozema J, Chardonnens A, Tosserams M, Hafkenscheid R, Bruijnzeel S (1997) Leaf thickness and UV-B absorbing pigments of plants in relation to an elevational gradient along the Blue Mountains, Jamaica. Plant Ecology 128, 151–159.
Leaf thickness and UV-B absorbing pigments of plants in relation to an elevational gradient along the Blue Mountains, Jamaica.Crossref | GoogleScholarGoogle Scholar |

Ruhland CT, Day TA (1996) Changes in UV-B radiation screening effectiveness with leaf age in Rhododendron maximum. Plant, Cell & Environment 19, 740–746.
Changes in UV-B radiation screening effectiveness with leaf age in Rhododendron maximum.Crossref | GoogleScholarGoogle Scholar |

Savitch LV, Massacci A, Gray GR, Huner NPA (2000) Acclimation to low temperature or high light mitigates sensitivity to photoinhibition: roles of the Calvin cycle and the Mehler reaction. Australian Journal of Plant Physiology 27, 253–264.

Shen ZH, Liu ZL, Wu J (2004) Altitudinal pattern of flora on the eastern slope of Mt. Gongga. Biodiversity Science 12, 89–98. [in Chinese]

Shi Z, Liu S, Liu X, Centritto M (2006) Altitudinal variation in photosynthetic capacity, diffusional conductance and δ13C of butterfly bush (Buddleja davidii) plants growing at high elevations. Physiologia Plantarum 128, 722–731.
Altitudinal variation in photosynthetic capacity, diffusional conductance and δ13C of butterfly bush (Buddleja davidii) plants growing at high elevations.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXjslU%3D&md5=ef89754849c80ad0fc5005196fbc0282CAS |

Shipley B, Lechoweicz MJ, Wright IJ, Reich PB (2006) Fundamental trade-offs generating the worldwide leaf economics spectrum. Ecology 87, 535–541.
Fundamental trade-offs generating the worldwide leaf economics spectrum.Crossref | GoogleScholarGoogle Scholar | 16602282PubMed |

Skaltsa H, Verykokidou E, Harvala C, Karabourniotis G, Manetas Y (1994) UV-B protective potential and flavonoid content of leaf hairs of Quercus ilex. Phytochemistry 37, 987–990.
UV-B protective potential and flavonoid content of leaf hairs of Quercus ilex.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2MXitFyks74%3D&md5=10a11bee075af2ce194282424e175226CAS | 1:CAS:528:DyaK2MXitFyks74%3D&md5=10a11bee075af2ce194282424e175226CAS |

Sobrado MA (1997) Embolism vulnerability in drought-deciduous and evergreen species of a tropical dry forest. Acta Oecologica 18, 383–391.
Embolism vulnerability in drought-deciduous and evergreen species of a tropical dry forest.Crossref | GoogleScholarGoogle Scholar |

Takashima T, Hikosaka K, Hirose T (2004) Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species. Plant, Cell & Environment 27, 1047–1054.
Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXnt1Kks7o%3D&md5=3e8595dff235309c2d9e765cefc56385CAS | 1:CAS:528:DC%2BD2cXnt1Kks7o%3D&md5=3e8595dff235309c2d9e765cefc56385CAS |

Thomas A (1997) The climate of the Gongga Shan Range, Sichuan Province, PR China. Arctic and Alpine Research 29, 226–232.
The climate of the Gongga Shan Range, Sichuan Province, PR China.Crossref | GoogleScholarGoogle Scholar |

Thomas SC (2011) Genetic vs. phenotypic responses of trees to altitude. Tree Physiology 31, 1161–1163.
Genetic vs. phenotypic responses of trees to altitude.Crossref | GoogleScholarGoogle Scholar | 22084019PubMed |

Van Arendonk JJCM, Poorter H (1994) The chemical composition and anatomical structure of leaves of grass species differing in relative growth rate. Plant, Cell & Environment 17, 963–970.
The chemical composition and anatomical structure of leaves of grass species differing in relative growth rate.Crossref | GoogleScholarGoogle Scholar |

Wang L, Ouyang H, Zhou CP, Zhang F, Bai JH, Peng K (2004) Distribution characteristics of soil organic matter and nitrogen on the eastern slope of Mt. Gongga. Acta Geographica Sinica 59, 1012–1019.

Warton DI, Wright IJ, Falster DS, Westoby M (2006) Bivariate line-fitting methods for allometry. Biological Reviews of the Cambridge Philosophical Society 81, 259–291.
Bivariate line-fitting methods for allometry.Crossref | GoogleScholarGoogle Scholar | 16573844PubMed |

Webb CO, Ackerly DD, Kembel SW (2008) Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Bioinformatics 24, 2098–2100.
Phylocom: software for the analysis of phylogenetic community structure and trait evolution.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXhtFWhurjM&md5=c7dab9dd69278ed1993cb080b42e54cbCAS | 1:CAS:528:DC%2BD1cXhtFWhurjM&md5=c7dab9dd69278ed1993cb080b42e54cbCAS | 18678590PubMed |

Wellburn A (1994) The spectral determination of chlorophylls a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resoultion. Journal of Plant Physiology 144, 307–313.
The spectral determination of chlorophylls a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resoultion.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2cXmsFShsbY%3D&md5=f52ea987007a40e95d42cae7eeb856fbCAS | 1:CAS:528:DyaK2cXmsFShsbY%3D&md5=f52ea987007a40e95d42cae7eeb856fbCAS |

Westoby M (1998) A leaf-height-seed (LHS) plant ecology strategy scheme. Plant and Soil 199, 213–227.
A leaf-height-seed (LHS) plant ecology strategy scheme.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1cXjslOht7o%3D&md5=bcd0d28bdeca38153fed27f3dc45f148CAS | 1:CAS:528:DyaK1cXjslOht7o%3D&md5=bcd0d28bdeca38153fed27f3dc45f148CAS |

Westoby M, Falster D, Moles A, Vesk P, Wright IJ (2002) Plant ecological strategies: some leading dimensions of variation between species. Annual Review of Ecology Evolution and Systematics 33, 125–159.
Plant ecological strategies: some leading dimensions of variation between species.Crossref | GoogleScholarGoogle Scholar |

Woodward F (1979) The differential temperature responses of the growth of certain plant species from different altitudes. II. Analyses of the control and morphology of leaf extension and specific leaf area of Phleum bertolonii D.C. and P. alpinum L. New Phytologist 82, 397–405.
The differential temperature responses of the growth of certain plant species from different altitudes. II. Analyses of the control and morphology of leaf extension and specific leaf area of Phleum bertolonii D.C. and P. alpinum L.Crossref | GoogleScholarGoogle Scholar |

Wright IJ, Westoby M (2002) Leaves at low versus high rainfall: coordination of structure, lifespan and physiology. New Phytologist 155, 403–416.
Leaves at low versus high rainfall: coordination of structure, lifespan and physiology.Crossref | GoogleScholarGoogle Scholar |

Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets Ü, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004) The worldwide leaf economics spectrum. Nature 428, 821–827.
The worldwide leaf economics spectrum.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXjt1Crt74%3D&md5=ecab0c2b186489650165ad106cd5e1adCAS | 1:CAS:528:DC%2BD2cXjt1Crt74%3D&md5=ecab0c2b186489650165ad106cd5e1adCAS | 15103368PubMed |

Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Garnier E, Hikosaka K, Lamont BB, Lee W, Oleksyn J, Osada N, Poorter H, Villar R, Warton DI, Westoby M (2005) Assessing the generality of global leaf trait relationships. New Phytologist 166, 485–496.
Assessing the generality of global leaf trait relationships.Crossref | GoogleScholarGoogle Scholar | 15819912PubMed |

Yang DM, Li GY, Sun SC (2008) The generality of leaf size versus number tradeoff in temperate woody species. Annals of Botany 102, 623–629.
The generality of leaf size versus number tradeoff in temperate woody species.Crossref | GoogleScholarGoogle Scholar |

Zhang SB, Zhou ZK, Hu H, Xu K, Yan N, Li SY (2005) Photosynthetic performances of Quercus pannosa vary with altitude in the Hengduan Mountains, southwest China. Forest Ecology and Management 212, 291–301.
Photosynthetic performances of Quercus pannosa vary with altitude in the Hengduan Mountains, southwest China.Crossref | GoogleScholarGoogle Scholar |

Zhong X, Wu N, Luo J, Yin KP, Tang Y, Pan ZF (1997). ‘Researches of the forest ecosystems on Gongga Mountains.’ (Chengdu Science and Technology University Press, Chengdu) [in Chinese]

Zhong XH, Zhang WJ, Luo J (1999) The characteristics of the mountain ecosystem and environment in the Gongga Mountain region. Ambio 28, 648–654. [in Chinese]