Review papers

Structural attributes of stand overstory and light under the canopy



This paper reviews the literature relating to the relationship between light availability in the understory and the main qualitative and quantitative attributes of stand overstory usually considered in forest management and planning (species composition, density, tree sizes, etc.) as well as their changes as consequences of harvesting. The paper is divided in two sections: the first one reviews studies which investigated the influence of species composition on understory light conditions; the second part examines research on the relationships among stand parameters determined from dendrometric field data and the radiation on understory layer. The objective was to highlight which are the most significant stand traits and management features to build more practical models for predicting light regimes in any forest stand and, in more general terms, to support forest managers in planning and designing silvicultural treatments that retain structure in different way in order to meet different objectives.


Understory light; structural attributes; overstory stand; forest management

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Ameztegui A., Coll L., Benavides R., Valladares F., Paquette A. 2012 - Understory light predictions in mixed conifer mountain forests: Role of aspect-induced variation in crown geometry and openness. Forest Ecology and Management 276: 52-61. doi:10.1016/j.foreco.2012.03.021

Aschoff T., Thies M., Spiecker H. 2004 - Describing forest stands using terrestrial laser- scanning. In: Conference Proceedings ISPRS Conference. ISPRS International Archives of Photogrammetry, Remote Sensing and Spatial Information Sciences, Istanbul, Turkey, 12–23 July, vol. XXXV, Part B: 237–241.

Astrup R., Larson B. C. 2006 - Regional variability of species-specific crown openness for aspen and spruce in western boreal Canada. Forest Ecology and Management 228: 241-250. doi:10.1016/j.foreco.2006.02.048

Aussenac G. 2000 - Interactions between forest stands and microclimate: Ecophysiological aspects and consequences for silviculture. Annals of Forest Science 57: 287–301. doi: 10.1051/forest:2000119

Barbier S., Gosselin F., Balandier P. 2008 - Influence of tree species on understory vegetation diversity and mechanisms involved-A critical review for temperate and boreal forests. Forest Ecology and Management 254: 1–15. doi:10.1016/j.foreco.2007.09.038

Battaglia M.A., Mou P., Palik B., Mitchell R.J. 2002 - The effect of spatially variable overstory on the understory light environment of an open-canopied longleaf pine forest. Canadian Journal of Forest Research 32 (11): 1984-1991. doi:10.1139/x02-087

Bayer D., Seifert S., Pretzsch H. 2013 - Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning. Trees 27: 1035-1047. doi: 10.1007/s00468-013-0854-4

Beaudet M., Messier C., Canham C.D. 2002 - Predictions of understorey light conditions in northern hardwood forests following parameterization, sensitivity analysis, and tests of the SORTIE light model. Forest Ecology and Management 165: 235-248.

Beaudet M., Harvey B. D., Messier C., Coates K. D., Poulin J., Kneeshaw D. D., Brais S.,

Bergeron Y. 2011 - Managing understory light conditions in boreal mixedwoods through variation in the intensity and spatial pattern of harvest: A modelling approach. Forest ecology and Management 261: 84-94. doi: 10.1016/j.foreco.2010.09.033

Bond B. J., Farnsworth B. T., Coulombe R. A., Winner W. E. 1999 - Foliage physiology and biochemistry in response to light gradients in conifers with varying shade tolerance. Oecologia 120: 183-192. doi: 10.1007/s004420050847

Bradford J. B., Kastendick D. N. 2010 - Age-related patterns of forest complexity and carbon storage in pine and aspen-birch ecosystems of northern Minnesota, USA. Canadian Journal of Forest Research 40 (3): 401-409. doi: l 0.1 I 39/X I0-002

Canham C. D., Denslow J. S., Platt W. J., Runkle J. R., Spies T. A., White P. S. 1990 - Light regimes beneath closed canopies and tree-fall gaps in temperate and tropical forests. Canadian Journal of Forest Research 205: 620-631.

Canham C. D., Finzi A. C., Pacala S. W., Burbank D. H. 1994 - Causes and consequences of resource heterogeneity in forest: interspecific variation in light trasmission by canopy trees. Canadian Journal of Forest Research 24 (2): 337-349. doi: 10.1139/x94-046

Canhaman C., Coates K. D., Bartemucci P., Quaglia S. 1999 - Measurement and modeling of spatially explicit variation in light transmission through interior cedar-hemlock forests of British Columbia. Canadian Journal of Forest Research 29 (11): 1775-1783. doi: 10.1139/x99-151

Chianucci F., Cutini A. 2012 - Digital hemispherical photography for estimating forest canopy properties: current controversies and opportunities. iForest – Biogeosciences and Forestry 5: 290-295.

Chianucci F., Cutini A. 2013 - Estimation of canopy properties in deciduous forests with digital hemispherical and cover photography. Agricultural and Forest Meteorology 168: 130-1393.

Chianucci A., Cutini A., Corona P., Puletti N. 2014a - Estimation of leaf area index in understory deciduous trees using digital photography. Agricultural and Forest Meteorology 198: 259-264

Chianucci F., Macfarlane C., Pisek J., Cutini A., Casa R. 2014b - Estimation of foliage clumping from the LAI-2000 Plant Canopy Analyzer: effect of view caps. Trees (early view) doi. 10.1007/s00468-014-1115-x

Ciancio O. 2011 - Systemic silviculture: philosophical, epistemological and methodological aspects. L’Italia Forestale e Montana 66 (3): 181-190. doi: 10.4129/ifm.2011.3.01

Coates K. D., Canhaman C. D., Beaudet M., Sachs, D. L. Messier C. 2003 - Use of a spatially explicit individual-tree model (SORTIE/BC) to explore the implications of patchiness in structurally complex forests. Forest Ecology and Management 186: 297-310. doi:10.1016/S0378-1127(03)00301-3

Comeau P. G. 2001- Relationships between stand parameters and understorey light in boreal aspen stands. BC Journal of Ecosystems and Management 2: 1-8.

Comeau P. G., Heineman J. L. 2003 - Predicting understory light microclimate from stand parameters in young paper birch (Betula papyrifera Marsh.) stands. Forest Ecology and Management 180: 303-315. doi: 10.1016/S0378-1127(02)00581-9

Comeau P., Heineman J., Newsome T. 2006 - Evaluation of relationships between understory light and aspen basal area in the British Columbia central interior. Forest Ecology and Management 226: 80-87. doi:10.1016/j.foreco.2005.12.060

Danson F. M., Hetherington D., Morsdorf F., Koetz B., Allgöwer B. 2007 - Forest Canopy Gap Fraction From Terrestrial Laser Scanning. Geoscience and Remote Sensing Letters, IEEE 4: 157-160. doi: 10.1109/LGRS.2006.887064

Drever C. R., Lertzman K. P. 2001 - Light-growth responses of coastal Douglas-fir and western redcedar saplings under different regimes of soil moisture and nutrients. Canadian Journal of Forest Research 31 (12): 2124-2133. doi: 10.1139/cjfr-31-12-2124

Drever C. R., Lertzman K. P. 2003 - Effects of a wide gradient of retained tree structure on understory light in coastal Douglas-fir forests. Canadian Journal of Forest Research 33 (1): 137–146. doi: 10.1139/X02-167

Falster D. S., Westoby M. 2003 - Leaf size and angle vary widely across species: what consequences for light interception? New Phytologist 158: 509-525. doi: 10.1046/j.1469-8137.2003.00765.x

Franklin J. F., Spies T. A., Van Pelt R., Carey A. B., Thornburgh D. A., Berg D. R., Lindenmayer D. B., Harmon M. E., Keeton W. S., Shaw D. C., Ken B., Jiquan C. 2002 - Disturbances and structural development of natural forest ecosystems with silvicultural implications, using Douglas-fir forests as an example. Forest Ecology and Management 155: 399-423. doi:10.1016/S0378-1127(01)00575-8

Gamborg C., Larsen J. B. 2003 - ‘Back to nature’ - a sustainable future for forestry? Forest Ecology and Management 179: 559-571. doi: 10.1016/S0378-1127(02)00553-4

Geldenhuys C. J. 2010 - Managing forest complexity through application of disturbance- recovery knowledge in development of silvicultural systems and ecological rehabilitation in natural forest systems in Africa. Journal of Forest Research 15: 3-13. doi: 10.1007/s10310-009-0159-z

Gendron F., Messier C., Comeau P. G. 1998 - Comparison of various methods for estimating the mean growing season percent photosynthetic photon flux density in forests. Agricultural and Forest Meteorology 92: 55-70. doi:10.1016/S0168-1923(98)00082-3

Gendron F., Messier C., Comeau P.G. 2001 - Temporal variations in the understorey photosynthetic photon flux density of a deciduous stand: the effects of canopy development, solar elevation, and sky conditions. Agricultural and Forest Meteorology 106: 23-40. doi:10.1016/S0168-1923(00)00174-X

Gendron F., Messier C., Lo E., Comeau P. G. 2006 - The angular distribution of diffuse photosynthetically active radiation under different sky conditions in the open and within deciduous and conifer forest stands of Quebec and British Columbia, Canada. Annals of Forest Science 63: 43-53. doi: 10.1051/forest:2005096

Gilliam F. S. 2007 - The ecological significance of the herbaceous layer in temperate forest ecosystems. BioScience 57 (10): 845-858. doi: 10.1641/B571007

Grams T.E., Andersen C.P. 2007 - Competition for resources in trees: physiological versus

morphological plasticity. Progress in Botany 69: 356-381. doi: 10.1007/978-3-540-36832-8_16

Hale S.E. 2001 - Light regime beneath Sitka spruce plantations in northern Britain: preliminary results. Forest Ecology and Management 151: 61–66. doi:10.1016/S0378-1127(00)00696-4

Hale S. E. 2003 - The effect of thinning intensity on the below-canopy light environment in a Sitka spruce plantation. Forest Ecology and Management 179: 341-349. doi:10.1016/S0378-1127(02)00540-6

Hardy J.P., Melloh R., Koenig G., Marks D., Winstral A., Pomeroy J.W., Link T. 2004 - Solar radiation transmission through conifer canopies - Agricultural and Forest Meteorology 126: 257-270. doi:10.1016/j.agrformet.2004.06.012

Hart S. A., Chen H. Y. H. 2006 - Understory vegetation dynamics of North American boreal forests. Critical Reviews in Plant Sciences 25: 381-397. doi: 10.1080/07352680600819286

Heithecker T. D., Halpern C. B. 2007 - Edge-related gradients in microclimate in forest aggregates following structural retention harvests in western Washington. Forest Ecology and Management 248: 163-173. doi:10.1016/j.foreco.2007.05.003

Hikosaka K., Hirose T. 1997 - Leaf angle as a strategy for light competition: optimal and evolutionarily stable light-extinction coefficient within a leaf canopy. Ecoscience 4: 501-507.

Hunter J.C., Parker V.T., Barbour M.G. 1999 - Understory light and gap dynamics in an old-growth forested watershed in coastal California. Madrono 46: 1-6.

Jennings S.B., Brown N.D., Sheil D. 1999 - Assessing forest canopies and understorey illumination: canopy closure, canopy cover and other measures. Forestry 72 (1): 59–74. doi: 10.1093/forestry/72.1.59

Kato S., Komiyama A. 2002 - Spatial and seasonal heterogeneity in understory light conditions caused by differential leaf flushing of deciduous overstory trees. Ecological Research 17: 687-693. doi: 10.1046/j.1440-1703.2002.00529.x

Koch N. E., Skovsgaard J.P 1999 - Sustainable management of planted forests: some

comparisons between Central Europe and the United States. New Forests 17: 11–22. doi: 10.1023/A:1006520809425

Komiyama A., Kato S., Teranishi M. 2001a - Differential overstory leaf flushing contributes to the formation of patchy understory. Journal of Forest Research 6: 163–171. doi: 10.1007/BF02767088

Kuuluvainen T. 2009 - Forest Management and Biodiversity Conservation Based on Natural Ecosystem Dynamics in Northern Europe: The Complexity Challenge. A Journal of the Human Environment 38 (6): 309-315. doi:

Larcher W. 2003 - Physiological plant ecology – Ecophisiology and stress physiology of functional groups. 4th Edition. Springer, 513 p.

Larsen J.B., Nielsen A.B. 2007 - Nature-based forest management - Where are we going? Elaborating forest development types in and with practice. Forest Ecology and Management 238: 107–117. doi:10.1016/j.foreco.2006.09.087

Lefrançois M.L., Beaudet M., Messier C. 2008 - Crown openness as influenced by tree and site characteristics for yellow birch, sugar maple, and eastern hemlock. Canadian Journal of Forest Research 38 (3): 488-497. doi :10.1139/X07-177

Levin S. A. 1998 - Ecosystems and the biosphere as complex adaptive systems. Ecosystems 1: 431-436. doi: 10.1007/s100219900037

Lieffers V.J., Messier C., Stadt K.J., Gendron F., Comeau P.G. 1999 - Predicting and managing light in the understory of boreal forests. Canadian Journal of Forest Research 29 (6): 796-811. doi: 10.1139/x98-165

Maas H.G., Bienert A., Scheller S. Keane E. 2008 - Automatic forest inventory parameter determination from terrestrial laser scanner data. International Journal of Remote Sensing 29: 1579-1593. doi: 10.1080/01431160701736406

Macfarlane C., Hoffman M., Eamus D., Kerp N., Higginson S., McMurtrie R., Adams M.A. 2007 - Estimation of leaf area index in eucalypt forest using digital photography. Agricultural and Forest Meteorology 143: 176–188.

Malcom D. C., Mason W. L., Clarke G. C. 2001 - The transformation of conifer forests in Britain-regeneration, gap size and silvicultural systems - Forest Ecology and Management 151: 7-23. doi:10.1016/S0378-1127(00)00692-7

McKenzie D., Halpern C. B., Nelson C. R. 2000 - Overstory influences on herb and shrub communities in mature forests of western Washington, U.S.A. Canadian Journal of Forest Research 30 (10): 1655-1666. doi: 10.1139/x00-091

Meitner M. J., Sheppard S. R.J., Cavens D., Gandy R., Picard P., Harshaw H., Harrison D.

- The multiple roles of environmental data visualization in evaluating alternative forest management strategies. Computers and Electronics in Agriculture 49: 192-205. doi:10.1016/j.compag.2005.03.002

Messier C., Parent S., Bergeron Y. 1998 - Effects of overstory and understory vegetation on the understory light environment in mixed boreal forests. Journal of Vegetation Science 9: 511-520. doi: 10.2307/3237266

Messier C., Doucet R., Ruel J.C., Claveau Y., Kelly C., Lechowicz, M.J. 1999 - Functional ecology of advance regeneration in relation to light in boreal forests. Canadian Journal of Forest Research 29 (6): 812-823. doi: 10.1139/x99-070

Messier C., Puettmann K. 2011 - Forest as complex adaptive systems: implications for forest management and modelling. Italian Journal of Forest and Mountain Environments 66 (3): 249-258. doi: 10.4129/ifm.2011.3.11

Millar C. I., Stephenson N. L., Stephens S. L. 2007 - Climate change and forests of the future: managing in the face of uncertainty. Ecological Applications 17: 2145-2151.

Molder A., Bernhardt-Romermann M., Schmidt W. 2008 - Herb-layer diversity in deciduous forests: raised by tree richness or beaten by beech? Forest Ecology and Management 256: 272-281. doi:10.1016/j.foreco.2008.04.012

Monsi M., Saeki T. 2005 - On the factor light in plant communities and its importance for matter production. Annals of Botany 95: 549–567. doi:10.1093/aob/mci052

Montgomery R. A., Chadzon R. L. 2001 - Forest structure, canopy architecture, and light

transmittance in tropical wet forests. Ecology 82 (10): 2707-2718.

Nilson T. 1999 - Inversion of gap frequency data in forest stands. Agricoltural and Forest Meteorology 98-99: 437-448. doi:10.1016/S0168-1923(99)00114-8

Oker-Blom, P., Kellomäki S. 1982 - Theoretical computations on the role of crown shape in the absorption of light by forest trees. Mathematical Biosciences 59: 291-311.

O’Hara Kevin L. 1998 - Silviculture for structure diversity – a new look at multiaged systems. Journal of Forestry 96: 4-10.

O’Hara Kevin L., Ramage Benjamin S. 2013 - Silviculture in an uncertain world: utilizing multi-aged management systems to integrate disturbance. Forestry 86 (4): 401-410.doi: 10.1093/forestry/cpt012

Page L.M., Cameron A.D., Clarke G.C. 2001- Influence of overstorey basal area on density and growth of advance regeneration of Sitka spruce in variably thinned stands. Forest Ecology and Management 151: 25–35. doi:10.1016/S0378-1127(00)00693-9

Pisek J., Sonnentag O., Richardson A. D., Mõttus M. 2013 - Is the spherical leaf inclination

angle distribution a valid assumption for temperate and boreal broadleaf tree species? Agricultural and Forest Meteorology 169: 186–194. doi:10.1016/j.agrformet.2012.10.011

Pommerening A., Murphy S.T. 2004 - A review of the history, definitions and methods of continuous cover forestry with special attention to afforestation and restocking. Forestry 77 (1): 27-44. doi: 10.1093/forestry/77.1.27

Pretzsch H., Schütze G. 2009 - Transgressive overyielding in mixed compared with pure stands of Norway spruce and European beech in Central Europe: Evidence on stand level and explanation on individual tree level. European Journal of Forest Research 128: 183-204. doi: 10.1007/s10342-008-0215-9

Pretzsch H. 2014 - Canopy space filling and tree crown morphology in mixed-species stands compared with monocultures. Forest Ecology and Management 327: 251-264. doi:10.1016/j.foreco.2014.04.027

Pretzsch H., Schütze G. 2005 - Crown allometry and growing space efficiency of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) in pure and mixed stands. Plant Biology 7 (6): 628-639.

Promis A., Schindler D., Reif A., Cruz G. 2009 - Solar radiation transmission in and around canopy gaps in an uneven-aged Nothofagus betuloides forest. International Journal of Biometeorology 53: 355-367. doi:10.1007/s00484-009-0222-7

Puettmann K. J., Coates K. D., Messier C. 2009 - A critique of silviculture: managing for complexity. Island Press, Washington, D.C., USA, 189 p.

Puettmann K. 2011 - Silvicultural challenges and options in the context of global change: “simple” fixes and opportunities for new management approaches. Journal of Forestry 109 (6): 321-331.

Puettmann K. J., Messier C., Coates K. D. 2013 - Managing forests as complex adaptive systems: introductory concepts and applications. In: Messier C., Puettmann K. J., Coates K. D.: “Managing forests as complex adaptive systems. Building resilience to the challenge of global change”. Routledge, Abingdon: 3-16

Ross M. S., Flanagan L. B., La Roi G. H. 1986 - Seasonal and successional changes in light quality and quantity in the understory of boreal forest ecosystems. Canadian Journal of Botany 64 (11): 2792-2799. doi:10.1139/b86-373

Ross J., Sulev M. 2000 - Sources of errors in measurements of PAR. Agricultural and Forest Meteorology 100: 103-125. doi: 10.1016/S0168-1923(99)00144-6

Royo A. A., Carson W. P. 2006 - On the formation of dense understory layers in forests worldwide: consequences and implications for forest dynamics, biodiversity, and succession. Canadian Journal of Forest Research 36 (6): 1345-1362. doi: 10.1139/x06-025

Sonohat G., Balandier P., Ruchaud F. 2004 - Predicting solar radiation transmittance in the understory of even-aged coniferous stands in temperate forests. Annals of Forest Science 61 (7): 629-641. doi: 10.1051/forest:2004061

Sprugel D.G., Rascher K.G., Gersonde R., Dovciak M., Lutz J.A., Halpern C.B. 2009 - Spatially explicit modeling of overstory manipulations in young forests: effects on stand structure and light. Ecological Modelling 220: 3565-3575. doi:10.1016/j.ecolmodel.2009.07.029

Stenberg P., Kangas T., Smolander H., Linder S. 1999 - Shoot structure, canopy openness, and light interception in Norway spruce. Plant, Cell & Environment 22: 1133-1142. doi: 10.1046/j.1365-3040.1999.00484.x.

Thomas S. C., Halpern C. B., Falk D. A., Liguori D. A., Austin K. A. 1999 - Plant diversity in managed forests: understory responses to thinning and fertilization. Ecological Applications 9: 864-879.

Tinya F., Márialigeti S. , Király I. , Németh B. , Ódor P. 2009 - The effect of light conditions on herbs, bryophytes and seedlings of temperate mixed forests in Orség, Western Hungary. Plant Ecology 204: 69-81. doi: 10.1007/s11258-008-9566-z

Valladares F., Pearcy R.W. 1999 - The geometry of light interception by shoots of Heteromeles arbutifolia: morphological and physiological consequences for individual leaves. Oecologia 121:171-182. doi: 10.1007/s004420050919

Valladares F, Guzman B. 2006 - Canopy structure and spatial heterogeneity of understory light in an abandoned Holm oak woodland. Annals of Forest Science 63 (7): 749–761. doi:10.1051/forest:2006056

Wagner S., Nocentini S., Huth F., Hoogstra-Klein M. 2014 - Forest management approaches for coping with the Uncertainty of Climate Change: trade-offs in service provisioning and adaptability. Ecology and Society 19 (1): 32.

Waring R. H., Schlesinger W. H. 1985 - Forest ecosystems. Concepts and management. Academic Press London, 340 p.

Whigham D. F. 2004 - Ecology of woodland herbs in temperate deciduous forests. Annual Review of Ecology, Evolution and Systematics 35: 583-621. doi: 10.1139/x00-091

Yirdaw E., Luukkanen O. 2004 - Photosynthetically active radiation transmittance of forest plantation canopies in the Ethiopian highlands. Forest Ecology and Management 188: 17-24. doi:10.1016/j.foreco.2003.07.024

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