A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance
详细信息    查看全文
  • 作者:ülo Niinemets (1)
  • 关键词:Bifurcation ratio ; Biomass allocation ; Foliage distribution ; Leaf structure ; Light interception ; Review analysis ; Shade tolerance ; Shoot architecture ; Stand age
  • 刊名:Ecological Research
  • 出版年:2010
  • 出版时间:July 2010
  • 年:2010
  • 卷:25
  • 期:4
  • 页码:693-714
  • 全文大小:771KB
  • 参考文献:1. Aan A, Hallik L, Kull O (2006) Photon flux partitioning among species along a productivity gradient of an herbaceous plant community. J Ecol 94:1143-155
    2. Albaugh TJ, Allen HL, Dougherty PM, Kress LW, King JS (1998) Leaf area and above- and belowground growth responses of loblolly pine to nutrient and water additions. For Sci 44:317-28
    3. Ali MS, Kikuzawa K (2005) Plasticity in leaf-area density within the crown of / Aucuba japonica growing under different light levels. J Plant Res 118:307-16
    4. Anderson JM, Osmond CB (1987) Shade-sun response: compromises between acclimation and photoinhibition. In: Kyle DJ, Osmond CB, Arntzen CJ (eds) Photoinhibition. Elsevier, Amsterdam, pp?1-8
    5. Andersson F (1973) IBP-studies on plant productivity of south Swedish forest ecosystems. In: Kern L (ed) Modeling forest ecoystems, Report EDFB-IBP-73-7. Oak Ridge National Laboratory, Oak Ridge, pp?11-6
    6. Ando T (1981) IBP woodlands data set. In: Reichle DE (ed) Dynamic properties of forest ecosystems, IBP 23. Cambridge University Press, Cambridge, pp?604-05
    7. Anten NPR, Hirose T (1998) Biomass allocation and light partitioning among dominant and subordinate individuals in / Xanthium canadense stands. Ann Bot 82:665-73
    8. Baldocchi DD, Hutchison BA, Matt DR, McMillen RT (1985) Canopy radiative transfer models for spherical and known leaf inclination angle distributions: a test in an oak-hickory forest. J Appl Ecol 22:539-55
    9. Baltzer JL, Thomas SC (2007) Physiological and morphological correlates of whole-plant light compensation point in temperate deciduous tree seedlings. Oecologia 153:209-23
    10. Banez G, Gyokusen K, Saito A (1999) Plasticity in the branching characteristics of four year old / Quercus acutissima and / Q. serrata seedlings in response to low light intensity and additional fertilizer. Bull Kyushu Univ For 80:27-9
    11. Barclay HJ, Goodman D (2000) Conversion of total to projected leaf area index in conifers. Can J Bot 78:447-54
    12. Barker M, Van Miegroet H, Nicholas NS, Creed IF (2002) Variation in overstory nitrogen uptake in a small, high-elevation southern Appalachian spruce–fir watershed. Can J For Res 32:1741-752
    13. Bartelink HH (1998) A model of dry matter partitioning in trees. Tree Physiol 18:91-01
    14. Bassi R, Caffarri S (2000) Lhc proteins and the regulation of photosynthetic light harvesting function by xanthophylls. Photosynth Res 64:243-56
    15. Bégin C, Filion L (1999) Black spruce ( / Picea mariana) architecture. Can J Bot 77:664-72
    16. Bégué A, Hanan NP, Prince SD (1994) Radiative transfer in shrub savanna sites in Niger: preliminary results from HAPEX-Sahel. 2. Photosynthetically active radiation interception of the woody layer. Agric For Meteorol 69:247-66
    17. Bindiu C (1981) IBP woodlands data set. In: Reichle DE (ed) Dynamic properties of forest ecosystems, IBP 23. Cambridge University Press, Cambridge, p?614
    18. Bloor JMG, Grubb PJ (2004) Morphological plasticity of shade-tolerant tropical rainforest tree seedlings exposed to light changes. Funct Ecol 18:337-48
    19. Bond BJ, Farnsworth BT, Coulombe RA, Winner WE (1999) Foliage physiology and biochemistry in response to light gradients in conifers with varying shade tolerance. Oecologia 120:183-92
    20. Bond-Lamberty B, Wang C, Gower ST, Norman J (2002) Leaf area dynamics of a boreal black spruce fire chronosequence. Tree Physiol 22:993-001
    21. Borchert R, Slade NA (1981) Bifurcation ratios and the adaptive geometry of trees. Bot Gaz 142:394-01
    22. Borghetti M, Vendramin GG, Giannini R (1986) Specific leaf area and leaf area distribution in a young Douglas-fir plantation. Can J For Res 16:1283-288
    23. Cai Z-Q, Poorter L, Cao KF, Bongers F (2007) Seedling growth strategies in / Bauhinia species: comparing lianas and trees. Ann Bot 100:831-38
    24. Campbell GS (1986) Extinction coefficients for radiation in plant canopies calculated using an ellipsoidal inclination angle distribution. Agric For Meteorol 36:317-21
    25. Cescatti A (1997) Modelling the radiative transfer in discontinuous canopies of asymmetric crowns. II. Model testing and application in a Norway spruce stand. Ecol Model 101:275-84
    26. Cescatti A, Niinemets ü (2004) Sunlight capture. Leaf to landscape. In: Smith WK, Vogelmann TC, Chritchley C (eds) Photosynthetic adaptation: chloroplast to landscape. Springer, Berlin, pp?42-5
    27. Cescatti A, Zorer R (2003) Structural acclimation and radiation regime of silver fir ( / Abies alba Mill.) shoots along a light gradient. Plant Cell Environ 26:429-42
    28. Chen JM (1996) Optically-based methods for measuring seasonal variation of leaf area index in boreal conifer stands. Agric For Meteorol 80:135-63
    29. Chen HYH (1997) Interspecific responses of planted seedlings to light availability in interior British Columbia: survival, growth, allometric patterns, and specific leaf area. Can J For Res 27:1383-393
    30. Chen X, Vierling L, Deering D (2005) A simple and effective radiometric correction method to improve landscape change detection across sensors and across time. Remote Sens Environ 98:63-9
    31. Ciganda V, Gitelson A, Schepers J (2008) Vertical profile and temporal variation of chlorophyll in maize canopy: quantitative “crop vigor-indicator by means of reflectance-based techniques. Agron J 100:1409-417
    32. Classen AT, Chapman SK, Whitham TG, Hart SC, Koch GW (2007) Genetic-based plant resistance and susceptibility traits to herbivory influence needle and root litter nutrient dynamics. J Ecol 95:1181-194
    33. Claveau Y, Messier C, Comeau PG (2005) Interacting influence of light and size on aboveground biomass distribution in sub-boreal conifer saplings with contrasting shade tolerance. Tree Physiol 25:373-84
    34. Cordell S, Goldstein G, Meinzer FC, Vitousek PM (2001) Regulation of leaf life-span and nutrient-use efficiency of / Metrosideros polymorpha trees at two extremes of a long chronosequence in Hawaii. Oecologia 127:198-06
    35. Dahlberg U, Berge TW, Petersson H, Vencatasawmy CP (2004) Modelling biomass and leaf area index in a sub-arctic Scandinavian mountain area. Scand J For Res 19:60-1
    36. Day ME, Greenwood MS, White AS (2001) Age-related changes in foliar morphology and physiology in red spruce and their influence on declining photosynthetic rates and productivity with tree age. Tree Physiol 21:1195-204
    37. Deblonde G, Penner M, Royer A (1994) Measuring leaf area index with the LI-COR LAI-2000 in pine stands. Ecology 75:1507-511
    38. Delagrange S, Messier C, Lechowicz MJ, Dizengremel P (2004) Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability. Tree Physiol 24:775-84
    39. Delzon S, Loustau D (2005) Age-related decline in stand water use: sap flow and transpiration in a pine forest chronosequence. Agric For Meteorol 129:105-19
    40. Demmig-Adams B, Adams WW III (2006) Photoprotection in an ecological context: the remarkable complexity of thermal energy dissipation. Tansley review. New Phytol 172:11-1
    41. Diggle PK (1999) Heteroblasty and the evolution of flowering phenologies. Int J Plant Sci 160:S123–S134
    42. Duursma RA, M?kel? A (2007) Summary models for light interception and light-use efficiency of non-homogeneous canopies. Tree Physiol 27:859-70
    43. Ellsworth DS, Reich PB (1993) Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest. Oecologia 96:169-78
    44. Evans JR (1993) Photosynthetic acclimation and nitrogen partitioning within a lucerne canopy. I. Canopy characteristics. Aust J Plant Physiol 20:55-7
    45. Evans JR, Poorter H (2001) Photosynthetic acclimation of plants to growth irradiance: the relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain. Plant Cell Environ 24:755-67
    46. Falster DS, Westoby M (2003) Leaf size and angle vary widely across species: what consequences for light interception? New Phytol 158:509-25
    47. Farque L, Sinoquet H, Colin F (2001) Canopy structure and light interception in / Quercus petraea seedlings in relation to light regime and plant density. Tree Physiol 21:1257-267
    48. Fassnacht KS, Gower ST (1997) Interrelationships among the edaphic and stand characteristics, leaf area index, and aboveground net primary production of upland forest ecosystems in north central Wisconsin. Can J For Res 27:1058-067
    49. Fassnacht KS, Gower ST, Norman JM, McMurtrie RE (1994) A comparison of optical and direct methods for estimating foliage surface area index in forests. Agric For Meteorol 71:183-07
    50. Fisher JB (1986) Branching patterns and angles in trees. In: Givnish TJ (ed) On the economy of plant form and function. Proceedings of the sixth Maria Moors Cabot symposium. Evolutionary constraints on primary productivity: adaptive patterns of energy capture in plants. Harvard Forest, August 1983. Cambridge University Press, Cambridge, pp?493-23
    51. Fleck S (2003) Integrated analysis of relationships between 3D-structure, leaf photosynthesis and branch transpiration of mature / Fagus sylvatica and / Quercus petraea trees in a mixed forest stand. BIT?K, Bayreuth
    52. Fleck S, Niinemets ü, Cescatti A, Tenhunen JD (2003) Three-dimensional lamina architecture alters light harvesting efficiency in / Fagus: a leaf-scale analysis. Tree Physiol 23:577-89
    53. Fliervoet LM, Werger JA (1984) Canopy structure and microclimate of two wet grassland communities. New Phytol 96:115-30
    54. Frazer GW, Trofymow JA, Lertzman KP (2000) Canopy openness and leaf area in chronosequences of coastal temperate rainforests. Can J For Res 30:239-56
    55. Gersonde R, Battles JJ, O’Hara KL (2004) Characterizing the light environment in Sierra Nevada mixed-conifer forests using a spatially explicit light model. Can J For Res 34:1332-342
    56. Gholz HL (1982) Environmental limits on aboveground net primary production, leaf area, and biomass in vegetation zones of the Pacific Northwest. Ecology 63:469-81
    57. Gholz HL, Vogel SA, Cropper WP Jr, McKelvey K, Ewel KC, Teskey RO, Curran PJ (1991) Dynamics of canopy structure and light interception in / Pinus elliottii stands, North Florida. Ecol Monogr 61:33-1
    58. Givnish T (1979) On the adaptive significance of leaf form. In: Solbrig OT, Jain S, Johnson GB, Raven PH (eds) Topics in plant population biology. Columbia University Press, New York, pp?375-07
    59. Givnish TJ (1982) On the adaptive significance of leaf height in forest herbs. Am Nat 120:353-81
    60. Givnish TJ (1988) Adaptation to sun and shade: a whole-plant perspective. Aust J Plant Physiol 15:63-2
    61. Givnish TJ (2002) Adaptive significance of evergreen versus deciduous leaves: solving the triple paradox. Silva Fenn 36:703-43
    62. Givnish TJ, Vermeij GJ (1976) Sizes and shapes of liana leaves. Am Nat 110:743-76
    63. Gordon AG (1976) Taxonomy and genetics of / Picea rubens and its relationship to / Picea mariana. Can J Bot 54:781-13
    64. Gould KS (1993) Leaf heteroblasty in / Pseudopanax crassifolius: functional significance of leaf morphology and anatomy. Ann Bot 71:61-0
    65. Gower ST, Vogt KA, Grier CC (1992) Carbon dynamics of rocky mountain Douglas-fir: influence of water and nutrient availability. Ecol Monogr 62:43-5
    66. Gower ST, Reich PB, Son Y (1993) Canopy dynamics and aboveground production of five tree species with different leaf longevities. Tree Physiol 12:327-45
    67. Gower ST, Vogel J, Norman JM, Kucharik CJ, Steele SJ, Stow TK (1997) Carbon distribution and aboveground net primary production in aspen, jack pine and black spruce stands in Saskatchewan and Manitoba. J Geophys Res 102:29029-9041
    68. Gracia CA (1988) Chlorophyll content and leaf energetics of plant communities in south-eastern Queensland (Australia) in relation to water availability. In: di Castri F, C Floret, S Rambal, J Roy (eds) Time scales and water stress. Proceedings of the 5th International Conference on Mediterranean Ecosystems. International Union of Biological Sciences, Paris, pp?347-60
    69. Grassi G, Bagnaresi U (2001) Foliar morphological and physiological plasticity in / Picea abies and / Abies alba saplings along a natural light gradient. Tree Physiol 21:959-67
    70. Gratzer G, Darabant A, Chhetri PB, Rai PB, Eckmüllner O (2004) Interspecific variation in the response of growth, crown morphology, and survivorship to light of six tree species in the conifer belt of the Bhutan Himalayas. Can J For Res 34:1093-107
    71. Greenway KJ, Macdonald SE, Lieffers VJ (1992) Is long-lived foliage in / Picea mariana an adaptation to nutrient-poor conditions? Oecologia 91:184-91
    72. Greenwood MS (1995) Juvenility and maturation in conifers: current concepts. Tree Physiol 15:433-38
    73. Greenwood MS, Day ME, Berlyn GP (2009) Regulation of foliar plasticity in conifers: developmental and environmental factors. J Sustain For 28:48-2
    74. Grier CC, Running SW (1977) Leaf area of mature northwestern coniferous forests: relation to site water balance. Ecology 58:893-99
    75. Grier CC, Elliott KJ, McCullough DG (1992) Biomass distribution and productivity of / Pinus edulis-em class="a-plus-plus">Juniperus monosperma woodlands of north-central Arizona. For Ecol Manag 50:331-50
    76. Groninger JW, Seiler JR, Peterson JA, Kreh RE (1996) Growth and photosynthetic responses of four Virginia Piedmont tree species to shade. Tree Physiol 16:773-78
    77. Gutschick VP, Wiegel FW (1988) Optimizing the canopy photosynthetic rate by patterns of investment in specific leaf mass. Am Nat 132:67-6
    78. Hallik L, Kull O, Niinemets ü, Aan A (2009a) Contrasting correlation networks between leaf structure, nitrogen and chlorophyll in herbaceous and woody canopies. Basic Appl Ecol 10:309-18
    79. Hallik L, Niinemets ü, Wright IJ (2009b) Tolerance to shade and drought in relation to foliage structural and functional traits in temperate woody species. New Phytol 184:257-74
    80. Hansen U, Fiedler B, Rank B (2002) Variation of pigment composition and antioxidative systems along the canopy light gradient in a mixed beech/oak forest: a comparative study on deciduous tree species differing in shade tolerance. Trees Struct Funct 16:354-64
    81. Harcombe PA, Bill CJ, Fulton M, Glitzenstein JS, Marks PL, Elsik IS (2002) Stand dynamics over 18?years in a southern mixed hardwood forest, Texas, USA. J Ecol 90:947-57
    82. Harris JR, Bassuk NL (1993) Adaptation of trees to low light environments: effect on branching pattern of / Fraxinus americana. J Arboric 19:339-43
    83. Hashimoto R (1991) Canopy development in young sugi ( / Cryptomeria japonica) stands in relation to changes with age in crown morphology and structure. Tree Physiol 8:129-43
    84. Hay JA, Dellow UV (1952) New Zealand conifers: a note on their uses and importance. Tuatara 4:109-18
    85. Hemmerlein MT, Smith WK (1994) Structural scaling of light interception efficiency in / Picea engelmannii and / Abies lasiocarpa. Tree Physiol 14:1139-148
    86. Hessl AE, Milesi C, White MA, Peterson DL, Keane RE (2004) Ecophysiological parameters for Pacific Northwest trees. US Department of Agriculture, Forest Service, Pacific Northwest Research Station, Portland, OR
    87. Hikosaka K, Hirose T (1997) Leaf angle as a strategy for light competition: optimal and evolutionary stable light extinction coefficient within a leaf canopy. écoscience 4:501-07
    88. Hikosaka K, Terashima I (1995) A model of the acclimation of photosynthesis in the leaves of C3 plants to sun and shade with respect to nitrogen use. Plant Cell Environ 18:605-18
    89. Hikosaka K, Terashima I (1996) Nitrogen partitioning among photosynthetic components and its consequence in sun and shade plants. Funct Ecol 10:335-43
    90. Hirose T, Werger MJA (1995) Canopy structure and photon flux partitioning among species in a herbaceous plant community. Ecology 76:466-74
    91. Holder CD (2000) Geography of / Pinus elliottii Engelm and / Pinus palustris Mill. leaf life-spans in the southeastern USA. J Biogeogr 27:311-18
    92. Honda H, Fisher JB (1978) Tree branch angle: maximizing effective leaf area. Science 199:888-90
    93. Horn HS (1971) The adaptive geometry of trees. Princeton University Press, Princeton
    94. Huang J, Wu Z, Guo SR (2007) Efects of low light intensity on photosynthetic capacity, photosynthetic nitrogen utilization efficiency and nitrogen partitioning in non-heading Chinese cabbage leaves. Acta Hortic 761:235-42
    95. Iio A, Fukasawa H, Nose Y, Kato S, Kakubari Y (2005) Vertical, horizontal and azimuthal variations in leaf photosynthetic characteristics within a / Fagus crenata crown in relation to light acclimation. Tree Physiol 25:525-36
    96. Ishii H, Asano S (2009) The role of crown architecture, leaf phenology and photosynthetic activity in promoting complementary use of light among coexisting species in temperate forests. Ecol Res doi:10.1007/s11284-009-0668-4
    97. Iwasa Y, Cohen D, Leon JA (1984) Tree height and crown shape, as a result of competitive games. J Theor Biol 112:279-97
    98. Jack SB, Long JN (1992) Forest production and the organization of foliage within crowns and canopies. For Ecol Manag 49:233-45
    99. James SA, Bell DT (1996) Leaf orientation of juvenile / Eucalyptus camaldulensis. Aust J Bot 44:139-56
    100. James SA, Bell DT (2000) Leaf orientation, light interception and stomatal conductance of / Eucalyptus globulus ssp / globulus leaves. Tree Physiol 20:815-23
    101. Jarvis PG, Leverenz JW (1983) Productivity of temperate, deciduous and evergreen forests. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Physiological plant ecology. Springer, Berlin, pp?233-80
    102. Jenkins JC, Chojnacky DC, Heath LS, Birdsey RA (2004) Comprehensive database of diameter-based biomass regressions for North American tree species. US Department of Agriculture Forest Service, Northeastern Research Station, Newtown Square
    103. Kajimoto T, Matsuura Y, Osawa A, Abaimov AP, Zyryanova OA, Isaev AP, Yefremov DP, Mori S, Koike T (2006) Size–mass allometry and biomass allocation of two larch species growing on the continuous permafrost region in Siberia. For Ecol Manag 222:314-25
    104. Kawamura K (2009) A conceptual framework for the study of modular responses to local environmental heterogeneity within the plant crown and a review of related concepts and ideas. Ecol Res doi:10.1007/s11284-009-0688-0
    105. Kayama M, Sasa K, Koike T (2002) Needle life span, photosynthetic rate and nutrient concentration of / Picea glehnii, / P jezoensis and / P. abies planted on serpentine soil in northern Japan. Tree Physiol 22:707-16
    106. Kempf JS, Pickett STA (1981) The role of branch length and angle in branching pattern of forest shrubs along a successional gradient. New Phytol 88:111-16
    107. Kennedy MC (2009) Functional–structural models optimize the placement of foliage units for multiple whole-canopy functions. Ecol Res doi:10.1007/s11284-009-0658-6
    108. Kikuzawa K (2003) Phenological and morphological adaptations to the light environment in two woody and two herbaceous plant species. Funct Ecol 17:29-8
    109. King DA (2003) Allocation of above-ground growth is related to light in temperate deciduous saplings. Funct Ecol 17:482-88
    110. Kiniry JR (1998) Biomass accumulation and radiation use efficiency of honey mesquite and eastern red cedar. Biomass Bioenergy 15:467-73
    111. Kitajima K (1994) Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees. Oecologia 98:419-28
    112. Kohyama T (1980) Growth pattern of / Abies mariesii saplings under conditions of open-growth and suppression. Bot Mag Tokyo 93:13-4
    113. Kubota Y (2006) Spatial pattern and regeneration dynamics in a temperate / Abies-em class="a-plus-plus">Tsuga forest in southwestern Japan. J For Res 11:191-01
    114. Kull O, Aan A (1997) The relative share of graminoid and forb life-forms in a natural gradient of herb layer productivity. Ecography 20:146-54
    115. Kull O, Niinemets ü (1993) Variation in leaf morphometry and nitrogen concentration in / Betula pendula Roth., / Corylus avellana L. and / Lonicera xylosteum L. Tree Physiol 12:311-18
    116. Kull O, Tulva I (2000) Modelling canopy growth and steady-state leaf area index in an aspen stand. Ann For Sci 57:611-21
    117. Kull O, Koppel A, Noormets A (1998) Seasonal changes in leaf nitrogen pools in two / Salix species. Tree Physiol 18:45-1
    118. Kull O, Broadmeadow M, Kruijt B, Meir P (1999) Light distribution and foliage structure in an oak canopy. Trees Struct Funct 14:55-4
    119. Küppers M (1994) Canopy gaps: competitive light interception and economic space filling—a matter of whole-plant allocation. In: Caldwell MM, Pearcy RW (eds) Exploitation of environmental heterogeneity by plants. Ecophysiological processes above- and belowground. Academic, San Diego, pp?111-44
    120. Lagergren A, Eklundh L, Grelle A, Lundblad M, M?lder M, Lankreijer H, Lindroth A (2004) Net primary production and light use efficiency in a mixed coniferous forest in Sweden. Plant Cell Environ 28:412-23
    121. Le Roux X, Gauthier H, Bégué A, Sinoquet H (1997) Radiation absorption and use by humid savanna grassland: assessment using remote sensing and modelling. Agric For Meteorol 85:117-32
    122. Lee DW, Richards JH (1991) Heteroblastic development in vines. In: Mooney HA, Putz FH (eds) The biology of vines. Cambridge University Press, New York, pp?205-43
    123. Leverenz JW (1992) Shade shoot structure and productivity of evergreen conifer stands. Scand J For Res 7:345-53
    124. Leverenz JW, Hinckley TM (1990) Shoot structure, leaf area index and productivity of evergreen conifer stands. Tree Physiol 6:135-49
    125. Leverenz JW, Whitehead D, Stewart GH (2000) Quantitative analyses of shade-shoot architecture of conifers native to New Zealand. Trees Struct Funct 15:42-9
    126. Liang N, Nakadai T, Hirano T, Qu L, Koike T, Fujinuma Y, Inoue G (2004) In situ comparison of four approaches to estimating soil CO2 efflux in a northern larch ( / Larix kaempferi Sarg.) forest. Agric For Meteorol 123:97-17
    127. López-Serrano FR, Landete-Castillejos T, Martínez-Millán J, del Cerro-Barja A (2000) LAI estimation of natural pine forest using a non-standard sampling technique. Agric For Meteorol 101:95-11
    128. Lusk CH (2004) Leaf area and growth of juvenile temperate evergreens in low light: species of contrasting shade tolerance change rank during ontogeny. Funct Ecol 18:820-28
    129. Lusk CH, Contreras O (1999) Foliage area and crown nitrogen turnover in temperate rain forest juvenile trees of differing shade tolerance. J Ecol 87:973-83
    130. Lusk CH, Warton DI (2007) Global meta-analysis shows that relationships of leaf mass per area with species shade tolerance depend on leaf habit and ontogeny. New Phytol 176:764-74
    131. Lusk CH, Falster DS, Jara-Vergara CK, Jimenez-Castillo M, Salda?a-Mendoza A (2008a) Ontogenetic variation in light requirements of juvenile rainforest evergreens. Funct Ecol 22:454-59
    132. Lusk CH, Reich PB, Montgomery RA, Ackerly DD, Cavender-Bares J (2008b) Why are evergreen leaves so contrary about shade? Trends Ecol Evol 23:299-03
    133. Magnussen S, Smith VG, Yeatman CW (1986) Foliage and canopy characteristics in relation to aboveground dry matter increment of seven jack pine provenances. Can J For Res 16:464-70
    134. Marshall JD, Waring RH (1986) Comparison of methods of estimating leaf-area index in old-growth Douglas-fir. Ecology 67:975-79
    135. Mathieu F (2006) évolution de la transpiration après coupe dans la sapinière à bouleau blanc, Forêt Montmorency, Québec. In: Ma?trise en sciences forestières. Université Laval, Laval. http://archimede.bibl.ulaval.ca/archimede/files/d6feadca-d29f-24a81-b27a-26d24cdada28dea/23817.html
    136. McLeod SD, Running SW (1988) Comparing site quality indices and productivity in ponderosa pine stands of western Montana. Can J For Res 18:346-52
    137. McMillen GG, McClendon JH (1979) Leaf angle: an adaptive feature of sun and shade leaves. Bot Gaz 140:437-42
    138. Meir P, Kruijt B, Broadmeadow M, Barbosa E, Kull O, Carswell F, Nobre A, Jarvis PG (2002) Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area. Plant Cell Environ 25:343-57
    139. Mencuccini M, Grace J (1996) Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity. Tree Physiol 16:459-68
    140. Metcalfe DJ, Grubb PJ, Turner IM (1998) The ecology of very small-seeded shade-tolerant trees and shrubs in lowland rain forest in Singapore. Plant Ecol 134:131-49
    141. Miller PM, Eddleman LE, Miller JM (1995) / Juniperus occidentalis juvenile foliage: advantages and disadvantages for a stress-tolerant, invasive conifer. Can J For Res 25:470-79
    142. M?ller CM (1946) Untersuchungen über Laubmenge, Stoffverlust und Stoffproduktion des Waldes. Forstwiss Forsch 17:1-87
    143. Montague TG, Givnish TJ (1996) Distribution of black spruce versus eastern larch along peatland gradients: relationship to relative stature, growth rate, and shade tolerance. Can J Bot 74:1514-532
    144. Mori A, Takeda H (2004) Functional relationships between crown morphology and within-crown characteristics of understory saplings of three codominant conifers in a subalpine forest in central Japan. Tree Physiol 24:661-70
    145. Mori A, Mizumachi E, Sprugel DG (2008) Morphological acclimation to understory environments in / Abies amabilis, a shade- and snow-tolerant conifer species of the Cascade Mountains, Washington, USA. Tree Physiol 28:815-24
    146. Mugasha AG, Pluth DJ, Higginbotham KO, Takyi SK (1991) Foliar responses of black spruce to thinning and fertilization on a drained shallow peat. Can J For Res 21:152-63
    147. Myers BJ, Theiveyanathan S, O’Brien ND, Bond WJ (1996) Growth and water use of / Eucalyptus grandis and / Pinus radiata plantations irrigated with effluent. Tree Physiol 16:211-19
    148. Nel EM, Wessman CA (1993) Canopy transmittance models for estimating forest leaf area index. Can J For Res 23:2579-586
    149. Niinemets ü (1996) Changes in foliage distribution with tree size and relative irradiance: differences between the saplings of / Acer platanoides and / Quercus robur. Ecol Res 11:269-81
    150. Niinemets ü (1997a) Distribution patterns of foliar carbon and nitrogen as affected by tree dimensions and relative light conditions in the canopy of / Picea abies. Trees Struct Funct 11:144-54
    151. Niinemets ü (1997b) Role of foliar nitrogen in light harvesting and shade tolerance of four temperate deciduous woody species. Funct Ecol 11:518-31
    152. Niinemets ü (1998a) Adjustment of foliage structure and function to a canopy light gradient in two co-existing deciduous trees. Variability in leaf inclination angles in relation to petiole morphology. Trees Struct Funct 12:446-51
    153. Niinemets ü (1998b) Growth of young trees of / Acer platanoides and / Quercus robur along a gap—understory continuum: interrelationships between allometry, biomass partitioning, nitrogen, and shade-tolerance. Int J Plant Sci 159:318-30
    154. Niinemets ü (2002) Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in / Picea abies and / Pinus sylvestris. Tree Physiol 22:515-35
    155. Niinemets ü (2004) Adaptive adjustments to light in foliage and whole-plant characteristics depend on relative age in the perennial herb / Leontodon hispidus. New Phytol 162:683-96
    156. Niinemets ü (2005) Key plant structural and allocation traits depend on relative age in the perennial herb / Pimpinella saxifraga. Ann Bot 96:323-30
    157. Niinemets ü (2006) The controversy over traits conferring shade-tolerance in trees: ontogenetic changes revisited. J Ecol 94:464-70
    158. Niinemets ü (2007) Photosynthesis and resource distribution through plant canopies. Plant Cell Environ 30:1052-071
    159. Niinemets ü, Anten NPR (2009) Packing photosynthesis machinery: from leaf to canopy. In: Laisk A, Nedbal L, Govindjee (eds) Photosynthesis in silico: understanding complexity from molecules to ecosystems. Springer, Berlin, pp?363-399
    160. Niinemets ü, Kull O (1995a) Effects of light availability and tree size on the architecture of assimilative surface in the canopy of / Picea abies: variation in needle morphology. Tree Physiol 15:307-15
    161. Niinemets ü, Kull O (1995b) Effects of light availability and tree size on the architecture of assimilative surface in the canopy of / Picea abies: variation in shoot structure. Tree Physiol 15:791-98
    162. Niinemets ü, Kull O (1998) Stoichiometry of foliar carbon constituents varies along light gradients in temperate woody canopies: implications for foliage morphological plasticity. Tree Physiol 18:467-79
    163. Niinemets ü, Lukjanova A (2003a) Needle longevity, shoot growth and branching frequency in relation to site fertility and within-canopy light conditions in / Pinus sylvestris. Ann For Sci 60:195-08
    164. Niinemets ü, Lukjanova A (2003b) Total foliar area and average leaf age may be more strongly associated with branching frequency than with leaf longevity in temperate conifers. New Phytol 158:75-9
    165. Niinemets ü, Tenhunen JD (1997) A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species / Acer saccharum. Plant Cell Environ 20:845-66
    166. Niinemets ü, Bilger W, Kull O, Tenhunen JD (1998a) Acclimation to high irradiance in temperate deciduous trees in the field: changes in xanthophyll cycle pool size and in photosynthetic capacity along a canopy light gradient. Plant Cell Environ 21:1205-218
    167. Niinemets ü, Kull O, Tenhunen JD (1998b) An analysis of light effects on foliar morphology, physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance. Tree Physiol 18:681-96
    168. Niinemets ü, Kull O, Tenhunen JD (1999) Variability in leaf morphology and chemical composition as a function of canopy light environment in co-existing trees. Int J Plant Sci 160:837-48
    169. Niinemets ü, Ellsworth DS, Lukjanova A, Tobias M (2001) Site fertility and the morphological and photosynthetic acclimation of / Pinus sylvestris needles to light. Tree Physiol 21:1231-244
    170. Niinemets ü, Cescatti A, Lukjanova A, Tobias M, Truus L (2002a) Modification of light-acclimation of / Pinus sylvestris shoot architecture by site fertility. Agric For Meteorol 111:121-40
    171. Niinemets ü, Ellsworth DS, Lukjanova A, Tobias M (2002b) Dependence of needle architecture and chemical composition on canopy light availability in three North American / Pinus species with contrasting needle length. Tree Physiol 22:747-61
    172. Niinemets ü, Valladares F, Ceulemans R (2003) Leaf-level phenotypic variability and plasticity of invasive / Rhododendron ponticum and non-invasive / Ilex aquifolium co-occurring at two contrasting European sites. Plant Cell Environ 26:941-56
    173. Niinemets ü, Cescatti A, Christian R (2004) Constraints on light interception efficiency due to shoot architecture in broad-leaved / Nothofagus species. Tree Physiol 24:617-30
    174. Niinemets ü, Cescatti A, Rodeghiero M, Tosens T (2005a) Leaf internal diffusion conductance limits photosynthesis more strongly in older leaves of Mediterranean evergreen broad-leaved species. Plant Cell Environ 28:1552-566
    175. Niinemets ü, Lukjanova A, Sparrrow AD, Turnbull MH (2005b) Light-acclimation of cladode photosynthetic potentials in / Casuarina glauca: trade-offs between physiological and structural investments. Funct Plant Biol 32:571-82
    176. Niinemets ü, Sparrow A, Cescatti A (2005c) Light capture efficiency decreases with increasing tree age and size in the southern hemisphere gymnosperm / Agathis australis. Trees Struct Funct 19:177-90
    177. Niinemets ü, Portsmuth A, Tobias M (2006a) Leaf size modifies support biomass distribution between stems, petioles and mid-ribs in temperate plants. New Phytol 171:91-04
    178. Niinemets ü, Tobias M, Cescatti A, Sparrrow AD (2006b) Size-dependent variation in shoot light-harvesting efficiency in shade-intolerant conifers. Int J Plant Sci 167:19-2
    179. Niklas KJ (1994) Plant allometry: the scaling of form and process. University of Chicago Press, Chicago
    180. Nilson T (1971) A theoretical analysis of the frequency of gaps in plant stands. Agric Meteorol 8:25-8
    181. Nock CA, Caspersen JP, Thomas SC (2008) Large ontogenetic declines in intra-crown leaf area index in two temperate deciduous tree species. Ecology 89:744-53
    182. Norman JM, Campbell GS (1989) Canopy structure. In: Pearcy RW, Ehleringer JR, Mooney HA, Rundel PW (eds) Plant physiological ecology. Field methods and instrumentation. Chapman and Hall, London, pp?301-25
    183. O’Connell BM, Kelty MJ (1994) Crown architecture of understory and open-grown white pine ( / Pinus strobus L.) saplings. Tree Physiol 14:89-02
    184. Oguchi R, Hikosaka K, Hirose T (2005) Leaf anatomy as a constraint for photosynthetic acclimation: differential responses in leaf anatomy to increasing growth irradiance among three deciduous trees. Plant Cell Environ 28:916-27
    185. Olff H (1992) Effects of light and nutrient availability on dry matter and N allocation in six successional grassland species. Testing for resource ratio effects. Oecologia 89:412-21
    186. Oren R, Waring RH, Stafford SG, Barrett JW (1987) Twenty-four years of ponderosa pine growth in relation to canopy leaf area and understory competition. For Sci 33:538-47
    187. Oren R, Phillips N, Ewers BE, Pataki DE, Megonigal JP (1999) Sap-flux-scaled transpiration responses to light, vapor pressure deficit, and leaf area reduction in a flooded / Taxodium distichum forest. Tree Physiol 19:337-47
    188. Osunkoya OO, Omar-Ali K, Amit N, Dayan J, Daud DS, Sheng TK (2007) Comparative height–crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo. Am J Bot 94:1951-962
    189. Ovington JD (1957) Dry matter production by / Pinus sylvestris L. Ann Bot 21:287-14
    190. Paquette A, Bouchard A, Cogliastro A (2007) Morphological plasticity in seedlings of three deciduous species under shelterwood under-planting management does not correspond to shade tolerance ranks. For Ecol Manag 241:278-87
    191. Parker GG, Davis MM, Chapotin SM (2002) Canopy light transmittance in Douglas-fir-western hemlock stands. Tree Physiol 22:147-57
    192. Paz H (2003) Root/shoot allocation and root architecture in seedlings: variation among forest sites, microhabitats, and ecological groups. Biotropica 35:318-32
    193. Pearcy RW, Sims DA (1994) Photosynthetic acclimation to changing light environments: scaling from the leaf to the whole plant. In: Caldwell MM, Pearcy RW (eds) Exploitation of environmental heterogeneity by plants. Ecophysiological processes above- and belowground. Academic, San Diego, pp?145-74
    194. Pearcy RW, Valladares F, Wright SJ, Lasso de Paulis E (2004) A functional analysis of the crown architecture of tropical forest / Psychotria species: do species vary in light capture efficiency and consequently in carbon gain and growth? Oecologia 139:163-77
    195. Peichl M, Arain MA (2007) Allometry and partitioning of above- and belowground tree biomass in an age-sequence of white pine forests. For Ecol Manag 253:68-0
    196. Peterson DL, Spanner MA, Running SW, Teuber KB (1987) Relationship of thematic mapper simulator data to leaf area index of temperate coniferous forests. Remote Sens Environ 22:323-41
    197. Pickett STA, Kempf JS (1980) Branching patterns in forest shrubs and understory trees in relation to habitat. New Phytol 86:219-28
    198. Pierce LL, Running SW (1988) Rapid estimation of coniferous leaf area index using a portable integrating radiometer. Ecology 69:1762-767
    199. Planchais I, Sinoquet H (1998) Foliage determinants of light interception in sunny and shaded branches of / Fagus sylvatica L. Agric For Meteorol 89:241-53
    200. Pons TL (1976) An ecophysiological study in the field layer of ash coppice. I. Field measurements. Acta Bot Neerl 25:401-16
    201. Pons TL (1977) An ecophysiological study in the field layer of ash coppice. II. Experiments with / Geum urbanum and / Cirsium palustre in different light intensities. Acta Bot Neerl 26:29-2
    202. Poorter L, Werger MJA (1999) Light environment, sapling architecture, and leaf display in six rain forest tree species. Am J Bot 86:1464-473
    203. Poorter L, Bongers F, Sterck F, W?l H (2000) Architecture of 53 rain forest tree species differing in adult stature and shade tolerance. Ecology 84(3):602-08
    204. Poorter L, Bongers L, Bongers F (2006) Architecture of 54 moist-forest tree species: traits, trade-offs, and functional groups. Ecology 87:1289-301
    205. Poorter H, Niinemets ü, Poorter L, Wright IJ, Villar R (2009) Tansely review. Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytol 182:565-88
    206. Portsmuth A, Niinemets ü (2007) Structural and physiological plasticity to light and nutrients in five temperate deciduous woody species of contrasting shade tolerance. Funct Ecol 21:61-7
    207. Prior LD, Eamus D, Bowman DMJS (2003) Leaf attributes in the seasonally dry tropics: a comparison of four habitats in northern Australia. Funct Ecol 17:504-15
    208. Pronk AA (2004) Irrigation and nitrogen use efficiency of / Thuja occidentalis grown on sandy soils. In: de Wit (CT) Graduate school for production ecology and resource conservation. Wageningen University, Wageningen
    209. Pronk TE, Schieving F, Anten NPR, Werger MJA (2007) Plants that differ in height investment can coexist if they are distributing non-uniformly within an area. Ecol Complex 4:182-91
    210. Quero JL, Villar R, Mara?on T, Zamora R, Poorter L (2007) Seed-mass effects in four Mediterranean / Quercus species (Fagaceae) growing in contrasting light environments. Am J Bot 94:1795-803
    211. Rambal S (2001) Productivity of Mediterranean-type ecosystems. In: Mooney HA, Saugier B, Roy J (eds) Terrestrial global productivity: past present and future. Academic, San Diego, pp?315-44
    212. Reich PB, Walters MB, Ellsworth DS (1992) Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecol Monogr 62:365-92
    213. Reich PB, Tjoelker MG, Walters MB, Vanderklein DW, Buschena C (1998) Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedlings of nine boreal tree species grown in high and low light. Funct Ecol 12:327-38
    214. Richardson AD, Berlyn GP, Ashton PMS, Thadani R, Cameron IR (2000) Foliar plasticity of hybrid spruce in relation to crown position and stand age. Can J Bot 78:305-17
    215. Richardson AD, Ashton PMS, Berlyn GP, McGroddy ME, Cameron IR (2001) Within-crown foliar plasticity of western hemlock, / Tsuga heterophylla, in relation to stand age. Ann Bot 88:1007-015
    216. Rijkers T, Pons TL, Bongers F (2000) The effect of tree height and light availability on photosynthetic leaf traits of four neotropical species differing in shade tolerance. Funct Ecol 14:77-6
    217. Rikhari HC, Palni LMS, Sharma S, Nandi SK (1998) Himalayan yew: stand structure, canopy damage, regeneration and conservation strategy. Environ Conserv 25:334-41
    218. Ross J (1981) The radiation regime and architecture of plant stands. Junk, The Hague
    219. Rousseaux MC, Hall AJ, Sanchez RA (2000) Basal leaf senescence in a sunflower ( / Helianthus annuus) canopy: responses to increased R/FR ratio. Physiol Plant 110:477-82
    220. Runyon J, Waring RH, Goward SN, Welles JM (1994) Environmental limits on net primary production and light-use efficiency across the Oregon transect. Ecol Appl 4:226-37
    221. Ryan MG, Binkley D, Fownes JH (1997) Age-related decline in forest productivity: pattern and process. Adv Ecol Res 27:213-62
    222. Ryan MG, Binkley D, Fownes JH, Giardina CP, Senock RS (2004) An experimental test of the causes of forest growth decline with stand age. Ecol Monogr 74:393-14
    223. Ryel RJ, Beyschlag W (1995) Benefits associated with steep foliage orientation in two tussock grasses of the American Intermountain West. A look at water-use-efficiency and photoinhibition. Flora 190:251-60
    224. Saito S, Sato T, Kominami Y, Nagamatsu D, Kuramoto S, Sakai T, Tabuchi R, Sakai A (2004) Modeling the vertical foliage distribution of an individual / Castanopsis cuspidata (Thunb.) Schottky, a dominant broad-leaved tree in Japanese warm-temperate forest. Trees Struct Funct 18:486-91
    225. Sampson DA, Allen HL (1995) Direct and indirect estimates of leaf area index (LAI) for lodgepole and loblolly pine stands. Trees Struct Funct 9:119-22
    226. Sánchez-Gómez D, Valladares F, Zavala MA (2006) Functional traits and plasticity underlying shade tolerance in seedlings of four Iberian forest tree species. Tree Physiol 26:1425-433
    227. Satoo T (1974) Primary production relations of a young stand of / Metasequoia glyptostroboides planted in Tokyo: materials for the studies of growth in forest stands. 13. Bull Tokyo Univ For 66:153-64
    228. Schulze ED (1982) Plant life forms and their carbon, water and nutrient relations. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Physiological plant ecology. Springer, Berlin, pp?616-76
    229. Schulze ED, Fuchs M, Fuchs MI (1977) Spacial distribution of photosynthetic capacity and performance in a mountain spruce forest of northern Germany. III. The significance of the evergreen habit. Oecologia 30:239-48
    230. Scurlock JMO, Asner GP, Gower ST (2001) Worldwide historical estimates of leaf area index, 1932-000. Oak Ridge National Laboratory, Oak Ridge, TN
    231. Selaya NG, Anten NPR, Ooomen RJ, Matthies M, Werger MJA (2007) Above-ground biomass investments and light interception of tropical forest trees and lianas early in succession. Ann Bot 99:141-51
    232. Selaya NG, Ooomen RJ, Netten JJC, Werger MJA, Anten NPR (2008) Biomass allocation and leaf life span in relation to light interception by tropical forest plants during the first years of secondary succession. J Ecol 96:1211-221
    233. Sellin A (2000) Estimating needle area from geometric measurements: application of different calculation methods to Norway spruce. Trees Struct Funct 14:215-22
    234. Sheil D, Salim A, Chave J, Vanclay J, Hawthorne WD (2006) Illumination–size relationships of 109 coexisting tropical forest tree species. J Ecol 94:494-07
    235. Shibuya M, Hasaba H, Yajima T, Takahashi K (2005) Effect of thinning on allometry and needle-age distribution of trees in natural / Abies stands of northern Japan. J For Res 10:15-0
    236. Smith T, Huston M (1989) A theory of the spatial and temporal dynamics of plant communities. Vegetatio 83:49-9
    237. Smolander H, Stenberg P, Linder S (1994) Dependence of light interception efficiency of Scots pine shoots on structural parameters. Tree Physiol 14:971-80
    238. Sprugel DG, Brooks JR, Hinckley TM (1996) Effects of light on shoot geometry and needle morphology in / Abies amabilis. Tree Physiol 16:91-8
    239. Steingraeber DA, Kascht LJ, Frank DH (1979) Variation of shoot morphology and bifurcation ratio in sugar maple ( / Acer saccharum) saplings. Am J Bot 66:441-45
    240. Stenberg P, Linder S, Smolander H, Flower-Ellis J (1994) Performance of the LAI-2000 plant canopy analyzer in estimating leaf area index of some Scots pine stands. Tree Physiol 14:981-95
    241. Stenberg P, Smolander H, Sprugel DG, Smolander S (1998) Shoot structure, light interception, and distribution of nitrogen in an / Abies amabilis canopy. Tree Physiol 18:759-67
    242. Stenberg P, Kangas T, Smolander H, Linder S (1999) Shoot structure, canopy openness, and light interception in Norway spruce. Plant Cell Environ 22:1133-142
    243. Stenberg P, Palmroth S, Bond BJ, Sprugel DG, Smolander H (2001) Shoot structure and photosynthetic efficiency along the light gradient in a Scots pine canopy. Tree Physiol 21:805-14
    244. Stephens GR, Ward JS (1992) Sixty years of natural change in unmanaged mixed hardwood forests. Connecticut Agricultural Experiment Station, New Haven, Bull 902
    245. Sterck FJ, Bongers F (2001) Crown development in tropical rain forest trees: patterns with tree height and light availability. J Ecol 89:1-3
    246. Sterck FJ, Schieving F (2007) 3-D growth patterns of trees: effects of carbon economy, meristem activity, and selection. Ecol Monogr 77:405-20
    247. Sumida A, Komiyama A (1997) Crown spread patterns for five deciduous broad-leaved woody species: ecological significance of the retention patterns of larger branches. Ann Bot 80:759-66
    248. Tadaki Y (1966) Some discussions on the leaf biomass of forest stands and trees. Bull Gov For Exp Stn 184:135-61
    249. Tadaki Y (1991) Productivity of coniferous forests in Japan. In: Nakagoshi N, Golley FB (eds) Coniferous forest ecology, from an international perspective. Symposium held at the International Congress of Ecology, Yokohama, Japan, 1990. SPB, The Hague, pp?109-19
    250. Tadaki Y, Kawasaki Y (1966) Studies on the production structure of forest. IX. Primary productivity of a young / Cryptomeria plantation with excessively high stand density. J Jpn For Soc 48:55-1
    251. Tadaki Y, Ogata N, Nagatomo Y (1965) The dry matter productivity in several stands of / Cryptomeria japonica in Kyushu. Bull Gov For Exp Stn Tokyo 173:45-6
    252. Tadaki Y, Ogata N, Nagatomo Y, Yoshida T (1966) Studies on the production structure of forest. X. Primary productivity of an unthinned 45-year old stand of / Chamaecyparis obtusa. J Jpn For Soc 48:387-93
    253. Tadaki Y, Ogata N, Nagatomo Y (1967) Studies on the production structure of forest. XI. Primary productivities of 28-year old plantations of / Cryptomeria of cuttings and seedlings origin. Bull Gov For Exp Stn Tokyo 199:47-5
    254. Tadaki Y, Itatiya K, Tochiaki K, Miyauchi H, Matsuda U (1970) Studies on the production of structure of forest. XVI. Primary productivity of / Abies veitchii forests in subalpine zone of Mt. Fuji. Bull Gov For Exp Stn Tokyo 229:1-2
    255. Takahashi K, Lechowicz MJ (2008) Do interspecific differences in sapling growth traits contribute to the co-dominance of / Acer saccharum and / Fagus grandifolia? Ann Bot 101:103-09
    256. Takahashi K, Seino T, Kohyama T (2005) Plastic changes of leaf mass per area and leaf nitrogen content in response to canopy openings in saplings of eight deciduous broad-leaved tree species. Ecol Res 20:17-3
    257. Terashima I, Miyazawa S-I, Hanba YT (2001) Why are sun leaves thicker than shade leaves?—Consideration based on analyses of CO2 diffusion in the leaf. J Plant Res 114:93-05
    258. Terashima I, Hanba YT, Tazoe Y, Vyas P, Yano S (2006) Irradiance and phenotype: comparative eco-development of sun and shade leaves in relation to photosynthetic CO2 diffusion. J Exp Bot 57:343-54
    259. Thomas SC, Winner WE (2000) A rotated ellipsoidal angle density function improves estimation of foliage inclination distributions in forest canopies. Agric For Meteorol 100:19-4
    260. Toledo-Aceves T, Swaine MD (2008) Biomass allocation and photosynthetic responses of lianas and pioneer tree seedlings to light. Acta Oecol 34:38-9
    261. Torquebiau EF (1988) Photosynthetically active radiation environment, patch dynamics and architecture in a tropical rainforest in Sumatra. Aust J Plant Physiol 15:327-42
    262. Utsugi H (1999) Angle distribution of foliage in individual / Chamaecyparis obtusa canopies and effect of angle on diffuse light penetration. Trees Struct Funct 14:1-
    263. Utsugi H, Araki M, Kawasaki T, Ishizuka M (2006) Vertical distributions of leaf area and inclination angle, and their relationship in a 46-year-old / Chamaecyparis obtusa stand. For Ecol Manag 225:104-12
    264. Valladares F, Niinemets ü (2007) The architecture of plant crowns: from design rules to light capture and performance. In: Pugnaire FI, Valladares F (eds) Handbook of functional plant ecology. CRC, Boca Raton, pp?101-49
    265. Valladares F, Niinemets ü (2008) Shade tolerance, a key plant feature of complex nature and consequences. Annu Rev Ecol Evol Syst 39:237-57
    266. Valladares F, Chico JM, Aranda I, Balaguer L, Dizengremel P, Manrique E, Dreyer E (2002) The greater seedling high-light tolerance of / Quercus robur over / Fagus sylvatica is linked to a greater physiological plasticity. Trees Struct Funct 16:395-03
    267. Vanninen P, Ylitalo H, Siev?nen R, M?kel? A (1996) Effects of age and site quality on the distribution of biomass in Scots pine ( / Pinus sylvestris L.). Trees Struct Funct 10:231-38
    268. Veres JS, Pickett STA (1982) Branching patterns of / Lindera benzoin beneath gaps and closed canopies. New Phytol 91:767-72
    269. Vertessy RA, Watson FGR, O’Sullivan SK (2001) Factors determining relations between stand age and catchment water balance in mountain ash forests. For Ecol Manag 143:13-6
    270. Vogel JG, Gower ST (1998) Carbon and nitrogen dynamics of boreal jack pine stands with and without a green alder understory. Ecosystems 1:386-00
    271. Vose JM, Swank WT (1990) Assessing seasonal leaf area dynamics and vertical leaf area distribution in eastern white pine ( / Pinus strobus L.) with a portable light meter. Tree Physiol 7:125-34
    272. Walters MB, Reich PB (1999) Research review. Low-light carbon balance and shade tolerance in the seedlings of woody plants: do winter deciduous and broad-leaved evergreen species differ? New Phytol 143:143-54
    273. Walters MB, Kruger EL, Reich PB (1993) Growth, biomass distribution and CO2 exchange of northern hardwood seedlings in high and low light: relationships with successional status and shade tolerance. Oecologia 94:7-6
    274. Wang W-M, Li Z-L, Su H-B (2007) Comparison of leaf angle distribution functions: effects on extinction coefficient and fraction of sunlit foliage. Agric For Meteorol 143:106-22
    275. Ward JS, Stephens GR (1993) Influence of crown class and shade tolerance on individual tree development during deciduous forest succession in Connecticut, USA. For Ecol Manag 60:207-36
    276. Werger MJA, Hirose T (1988) Effects of light climate and nitrogen partitioning on the canopy structure of stands of a dicotyledonous, herbaceous vegetation. In: Werger MJA, van der Aart PJM, During HJ, Verhoeven JTA (eds) Plant form and vegetation structure adaptation plasticity and relation to herbivory. SPB, The Hague, pp?171-81
    277. Whitney GG (1976) The bifurcation ratio as an indicator of adaptive strategy in woody plant species. Bull Torrey Bot Club 103:67-2
    278. Whittaker RH (1966) Forest dimensions and production in the Great Smoky Mountains. Ecology 44:103-21
    279. Witkowski ETF, Lamont BB (1991) Leaf specific mass confounds leaf density and thickness. Oecologia 88:486-93
    280. 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 E, Villar R (2004) The world-wide leaf economics spectrum. Nature 428:821-27
    281. Xu M, Harrington TB (1998) Foliage biomass distribution of loblolly pine as affected by tree dominance, crown size, and stand characteristics. Can J For Res 28:887-92
  • 作者单位:ülo Niinemets (1)

    1. Institute of Agricultural and Environmental Sciences, Estonian University of Life Sciences, Kreutzwaldi 1, 51014, Tartu, Estonia
文摘
Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F L) modify light harvesting significantly. F L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M A, reductions in F L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.

© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号

地址:北京市海淀区学院路29号 邮编:100083

电话:办公室:(+86 10)66554848;文献借阅、咨询服务、科技查新:66554700