Effects of neighboring woody plants on target trees with emphasis on effects of understorey shrubs on overstorey physiology in forest communities: a mini-review
Abstract
Plant-plant interaction plays a key role in regulating the composition and structure of communities and ecosystems. Studies of plant-plant interactions in forest ecosystems have mainly concentrated on growth effects of neighboring plants on target trees. Physiological effects of neighboring plants on target trees, in particular understorey effects on physiology of overstorey trees, have received less attention. It is still unclear what is the physiological mechanisms underlying positive growth effects of understorey removal, although understorey removal has been applied to improve the wood production for hundreds of years worldwide. Only 17.5% of published works dealt with understorey-overstorey interactions and only a few of those researched the understorey effects on the physiology of overstorey trees. Case studies indicated that overstorey Abies faxoniana trees grown with different understorey shrubs showed significantly different levels of tissue nitrogen and mobile carbohydrates. Removal experiment showed that nitrogen and mobile carbohydrates concentrations in Cunninghamia lanceolata trees grown in the absence of understorey shrubs differed significantly (pure stand > mixture) with those in trees grown in the presence of understorey shrubs, in particular during the dry season. This review highlighted that the neighboring woody plants affect C-and N-physiology in overstorey trees. These effects may be mainly resulted from underground competition for soil water rather than for other resources as the effects were more pronounced during the dry season. The present review suggests that positive effects of neighboring removal (e.g., understorey removal, thinning) on overstorey trees can be expected more rapidly and strongly in stressful area (e.g., low rainfall, nutrient-poor site) than in areas with optimal growth conditions. Hence, ecophysiology-based management strategies for dealing with neighboring plants in forest ecosystems should take into account: 1) site conditions, 2) timing, duration and frequency of management practices, and 3) species-specific properties and other aspects such as biodiversity conservation and soil erosion.
Keywords
Carbon balance Competition Forest plantation management Nitrogen Non-structural carbohydrates Overstorey-understorey interaction Tree-shrub interaction Understorey removalAbbreviation
- NSC
Non-Structural Carbohydrate
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References
- Adams, P.R., C.L. Beadle, N.J. Mendham and P.J. Smethurst. 2003. The impact of timing and duration of grass control on growth of a young Eucalyptus globulus Labill. plantation. New Forests 26: 147–165.CrossRefGoogle Scholar
- Aerts, R. 1999. Interspecific competition in natural plant communities: mechanisms, trade-offs and plant-soil feedbacks. J. Exp. Bot. 50: 29–37.CrossRefGoogle Scholar
- Aphalo, P.J., C.L. Ballare and A.L. Scopel. 1999. Plant-plant signalling, the shade-avoidance response and competition. J. Exp. Bot. 50: 1629–1634.CrossRefGoogle Scholar
- Balandier, P., C. Collet, J.H. Miller, P.E. Reynolds and S.M. Zedaker. 2006. Designing forest vegetation management strategies based on the mechanisms and dynamics of crop tree competition by neighbouring vegetation. Forestry 79: 3–27.CrossRefGoogle Scholar
- Barnes, P.W. and S. Archer. 1999. Tree-shrub interactions in a subtropical savanna parkland: Competition or facilitation? J. Veg. Sci. 10: 525–536.CrossRefGoogle Scholar
- Barot, S. 2004. Mechanisms promoting plant coexistence: can all the proposed processes be reconciled? Oikos 106: 185–192.CrossRefGoogle Scholar
- Bauhus, J., A.P. van Winden and A.B. Nicotra. 2004. Aboveground interactions and productivity in mixed-species plantations of Acacia mearnsii and Eucalyptus globulus. Can. J. Forest Res. 34: 686–694.CrossRefGoogle Scholar
- Bertness, M.D. and R. Callaway. 1994. Positive interactions in communities. Trends Ecol. Evol. 9: 191–193.CrossRefPubMedPubMedCentralGoogle Scholar
- Binkley, D. 1983. Ecosystem production in Douglas-fir plantations -Interaction of red alder and site fertility. Forest Ecol. Manage. 5: 215–227.CrossRefGoogle Scholar
- Binkley, D., K.A. Dunkin, D. DeBell and M.G. Ryan. 1992. Production and nutrient cycling in mixed plantations of Eucalyptus and Albizia in Hawaii. Forest Sci. 38: 393–408.Google Scholar
- Binkley, D., J.D. Lousier and K. Cromack. 1984. Ecosystem effects of Sitka alder in a Douglas-fir plantation. Forest Sci. 30: 26–35.Google Scholar
- Binkley, D., R. Senock, S. Bird and T.G. Cole. 2003. Twenty years of stand development in pure and mixed stands of Eucalyptus saligna and nitrogen-fixing Facaltaria moluccana. Forest Ecol. Manage. 182: 93–102.CrossRefGoogle Scholar
- Binkley, D., J.L. Stape and M.G. Ryan. 2004. Thinking about efficiency of resource use in forests. Forest Ecol. Manage. 193: 5–16.CrossRefGoogle Scholar
- Boyden, S., D. Binkley and R. Senock. 2005. Competition and facilitation between Eucalyptus and nitrogen-fixing Falcataria in relation to soil fertility. Ecology 86: 992–1001.CrossRefGoogle Scholar
- Bréda, N., A. Granier and G. Aussenac. 1995. Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus petraea (Matt) Liebl). Tree Physiol. 15: 295–306.CrossRefGoogle Scholar
- Bristow, M., J.K. Vanclay, L. Brooks and M. Hunt. 2006. Growth and species interactions of Eucalyptus pellita in a mixed and monoculture plantation in the humid tropics of north Queensland. Forest Ecol. Manage. 233: 285–294.CrossRefGoogle Scholar
- Brooker, R.W. 2006. Plant-plant interactions and environmental change. New Phytol. 171: 271–284.CrossRefGoogle Scholar
- Brooker, R.W. and T.V. Callaghan. 1998. The balance between positive and negative plant interactions and its relationship to environmental gradients: a model. Oikos 81: 196–207.CrossRefGoogle Scholar
- Brooker, R.W., F.T. Maestre, R.M. Callaway, C.L. Lortie, L.A. Cavieres, G. Kunstler, P. Liancourt, K. Tielborger, J.M.J. Travis, F. Anthelme, C. Armas, L. Coll, E. Corcket, S. Delzon, E. Forey, Z. Kikvidze, J. Olofsson, F. Pugnaire, C.L. Quiroz, P. Saccone, K. Schiffers, M. Seifan, B. Touzard and R. Michalet. 2008. Facilitation in plant communities: the past, the present, and the future. J. Ecol. 96: 18–34.CrossRefGoogle Scholar
- Brozek, S. 1990. Effect of soil changes caused by red alder (Alnus rubra) on biomass and nutrient status of Douglas fir (Pseudo-tsuga menziesii) seedlings. Can. J. Forest Res. 20: 1320–1325.CrossRefGoogle Scholar
- Cakmak, I., A. Inal, A. Gunes and F. Zhang. 2007. Peanut/maize intercropping induced changes in rhizosphere and nutrient concentrations in shoots. Plant Physiol. Bioch. 45: 350–356.CrossRefGoogle Scholar
- Callaway, R.M. and L.R. Walker. 1997. Competition and facilitation: A synthetic approach to interactions in plant communities. Ecology 78: 1958–1965.CrossRefGoogle Scholar
- Carlyle, J.C. and D.C. Malcolm. 1986a. Nitrogen availability beneath pure spruce and mixed larch + spruce stands growing on a deep peat. 1. Net N mineralization measured by field and laboratory incubations. Plant and Soil 93: 95–113.CrossRefGoogle Scholar
- Carlyle, J.C. and D.C. Malcolm. 1986b. Nitrogen availability beneath pure spruce and mixed larch + spruce stands growing on a deep peat. 2. A comparison of N availability as measured by plant uptake and long-term laboratory incubations. Plant and Soil 93: 115–122.CrossRefGoogle Scholar
- Carter, G.A., J.H. Miller, D.E. Davis and R.M. Patterson. 1984. Effect of vegetative competition on the moisture and nutrient status of loblolly-pine. Can. J. Forest Res. 14: 1–9.CrossRefGoogle Scholar
- Casper, B.B. and R.B. Jackson. 1997. Plant competition underground. Annu. Rev. Ecol. Syst. 28: 545–570.CrossRefGoogle Scholar
- Curran, L.M., I. Caniago, G.D. Paoli, D. Astianti, M. Kusneti, M. Leighton, C.E. Nirarita and H. Haeruman. 1999. Impact of El Nino and logging on canopy tree recruitment in Borneo. Science 286: 2184–2188.CrossRefGoogle Scholar
- Davey, P.A., A.J. Parsons, L. Atkinson, K. Wadge and S.P. Long. 1999. Does photosynthetic acclimation to elevated CO2 increase photosynthetic nitrogen-use efficiency? A study of three native UK grassland species in open-top chambers. Funct. Ecol. 13: 21–28.CrossRefGoogle Scholar
- DeBell, D.S., T.G. Cole and C.D. Whitesell. 1997. Growth, development, and yield in pure and mixed stands of Eucalyptus and Albizia. Forest Sci. 43: 286–298.Google Scholar
- DeBell, D.S., C.D. Whitesell and T.H. Schubert. 1985. Mixed plantations of Eucalyptus and Leguminous trees enhance biomass production. USDA Forest Service Pacific Southwest Research Station Research Paper: 1–6.Google Scholar
- Devine, W.D. and T.B. Harrington. 2008. Belowground competition influences growth of natural regeneration in thinned Douglas-fir stands. Can. J. Forest Res. 38: 3085–3097.CrossRefGoogle Scholar
- Diaz-Espejo, A., E. Nicolas and J.E. Fernandez. 2007. Seasonal evolution of diffusional limitations and photosynthetic capacity in olive under drought. Plant Cell Environ. 30: 922–933.CrossRefGoogle Scholar
- Dudareva, N., F. Negre, D.A. Nagegowda and I. Orlova. 2006. Plant volatiles: Recent advances and future perspectives. Critical Rev. Plant Sci. 25: 417–440.CrossRefGoogle Scholar
- Elliott, K.J. and J.D. Knoepp. 2005. The effects of three regeneration harvest methods on plant diversity and soil characteristics in the southern Appalachians. Forest Ecol. Manage. 211: 296–317.CrossRefGoogle Scholar
- Elliott, K.J. and A.S. White. 1987. Competitive effects of various grasses and forbs on Ponderosa pine-seedlings. Forest Sci. 33: 356–366.Google Scholar
- Erskine, P.D., D. Lamb and M. Bristow. 2006. Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity? Forest Ecol. Manage. 233: 205–210.CrossRefGoogle Scholar
- Forrester, D.I., J. Bauhus and A.L. Cowie. 2005. Nutrient cycling in a mixed-species plantation of Eucalyptus globulus and Acacia mearnsii. Can. J. Forest Res. 35: 2942–2950.CrossRefGoogle Scholar
- Forrester, D.I., J. Bauhus and A.L. Cowie. 2006a. Carbon allocation in a mixed-species plantation of Eucalyptus globulus and Acacia mearnsii. Forest Ecol. Manage. 233: 275–284.CrossRefGoogle Scholar
- Forrester, D.I., J. Bauhus, A.L. Cowie and J.K. Vanclay. 2006b. Mixed-species plantations of Eucalyptus with nitrogen-fixing trees: A review. Forest Ecol. Manage. 233: 211–230.CrossRefGoogle Scholar
- Forrester, D.I., J. Bauhus, A.L. Cowie, P.A. Mitchell and J. Brockwell. 2007. Productivity of three young mixed-species plantations containing N2-fixing Acacia and non-N2-fixing Eucalyptus and Pinus trees in southeastern Australia. Forest Sci. 53: 426–434.Google Scholar
- Forrester, D.I., S. Theiveyanathan, J.J. Collopy and N.E. Marcar. 2010. Enhanced water use efficiency in a mixed Eucalyptus globulus and Acacia mearnsii plantation. Forest Ecol. Manage. 259: 1761–1770.CrossRefGoogle Scholar
- Fulbright, T.E., J.O. Kuti and A.R. Tipton. 1997. Effects of nurse-plant canopy light intensity on shrub seedling growth. J. Range Manage. 50: 607–610.CrossRefGoogle Scholar
- Gause, G.F. 1934. The Struggle for Existence. Williams and Wilkins, Baltimore.CrossRefGoogle Scholar
- Ginn, S.E., J.R. Seiler, B.H. Cazell and R.E. Kreh. 1991. Physiological and growth-responses of 8-year-old Loblolly-pine stands to thinning. Forest Sci. 37: 1030–1040.Google Scholar
- Grace, J.B. and D. Tilman. 1990. Perspectives on Plant Competition. Academic Press, New York.Google Scholar
- Grams, T.E.E., A.R. Kozovits, I.M. Reiter, J.B. Winkler, M. Som-merkorn, H. Blaschke, K.H. Haberle and R. Matyssek. 2002. Quantifying competitiveness in woody plants. Plant Biol. 4: 153–158.CrossRefGoogle Scholar
- Granhus, A. and F.H. Braekke. 2001. Nutrient status of Norway spruce stands subjected to different levels of overstorey removal. Trees-Structure and Function 15: 393–402.CrossRefGoogle Scholar
- Groninger, J.W., J.R. Seiler, J.A. Peterson and R.E. Kreh. 1996a. Growth and photosynthetic responses of four Virginia Piedmont tree species to shade. Tree Physiol. 16: 773–778.CrossRefGoogle Scholar
- Groninger, J.W., J.R. Seiler, S.M. Zedaker and P.C. Berrang. 1996b. Effects of CO2 concentration and water availability on growth and gas exchange in greenhouse-grown miniature stands of Loblolly Pine and Red Maple. Funct. Ecol. 10: 708–716.CrossRefGoogle Scholar
- Groninger, J.W., J.R. Seiler, S.M. Zedaker and P.C. Berrang. 1996c. Photosynthetic response of loblolly pine and sweetgum seedling stands to elevated carbon dioxide, water stress, and nitrogen level. Can. J. Forest Res. 26: 95–102.CrossRefGoogle Scholar
- Groninger, J.W., S.M. Zedaker and T.S. Fredericksen. 1997. Stand characteristics of inter-cropped loblolly pine and black locust. Forest Ecol. Manage. 91: 221–227.CrossRefGoogle Scholar
- Hagan, D.L., S. Jose, M. Thetford and K. Bohn. 2009. Production physiology of three native shrubs intercropped in a young long-leaf pine plantation. Agroforestry Systems 76: 283–294.CrossRefGoogle Scholar
- Harrington, C.A. and D.L. Reukema. 1983. Initial shock and longterm stand development following thinning in a Douglas-fir plantation. Forest Sci. 29: 33–46.Google Scholar
- Hart, S.A. and H.Y.H. Chen. 2006. Understory vegetation dynamics of North American boreal forests. Critical Rev. Plant Sci. 25: 381–397.CrossRefGoogle Scholar
- Hoch, G., M. Popp and C. Körner. 2002. Altitudinal increase of mobile carbon pools in Pinus cembra suggests sink limitation of growth at the Swiss treeline. Oikos 98: 361–374.CrossRefGoogle Scholar
- Hoch, G., A. Richter and C. Körner. 2003. Non-structural carbon compounds in temperate forest trees. Plant Cell Environ. 26: 1067–1081.CrossRefGoogle Scholar
- Hocker, H.W. 1982. Effects of thinning on biomass growth in young Populus tremuloides plots. Can. J. Forest Res. 12: 731–737.CrossRefGoogle Scholar
- Hokka, H., T. Penttila and B. Hanell. 1996. Effect of thinning on the foliar nutrient status of Scots pine stands on drained boreal peatlands. Can. J. Forest Res. 26: 1577–1584.CrossRefGoogle Scholar
- Jonard, M., L. Misson and Q. Ponette. 2006. Long-term thinning effects on the forest floor and the foliar nutrient status of Norway spruce stands in the Belgian Ardennes. Can. J. Forest Res. 36: 2684–2695.CrossRefGoogle Scholar
- Jose, S., R. Williams and D. Zamora. 2006. Belowground ecological interactions in mixed-species forest plantations. Forest Ecol. Manage. 233: 231–239.CrossRefGoogle Scholar
- Joyce, L.A. and R.L. Baker. 1987. Forest overstory understory relationships in Alabama forests. Forest Ecol. Manage. 18: 49–59.CrossRefGoogle Scholar
- Kaye, J.P., S.C. Resh, M.W. Kaye and R.A. Chimner. 2000. Nutrient and carbon dynamics in a replacement series of Eucalyptus and Albizia trees. Ecology 81: 3267–3273.CrossRefGoogle Scholar
- Keddy, P.A. 1989. Competition. Chapman and Hall, New York.CrossRefGoogle Scholar
- Kelty, M.J. 2006. The role of species mixtures in plantation forestry. Forest Ecol. Manage. 233: 195–204.CrossRefGoogle Scholar
- Kitajima, K. and K.P. Hogan. 2003. Increases of chlorophyll a/b ratios during acclimation of tropical woody seedlings to nitrogen limitation and high light. Plant Cell Environ. 26: 857–865.CrossRefGoogle Scholar
- Körner, C. 2003. Carbon limitation in trees. J. Ecol. 91: 4–17.CrossRefGoogle Scholar
- Kozovits, A.R., R. Matyssek, H. Blaschke, A. Gottlein and T.E.E. Grams. 2005. Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons. Global Change Biol. 11: 1387–1401.CrossRefGoogle Scholar
- Kume, A., T. Satomura, N. Tsuboi, M. Chiwa, Y.T. Hanba, K. Nakane, T. Horikoshi and H. Sakugawa. 2003. Effects of under-story vegetation on the ecophysiological characteristics of an overstory pine, Pinus densiflora. Forest Ecol. Manage. 176: 195–203.CrossRefGoogle Scholar
- Kuppers, B.I.L. 1996. Nitrogen and Rubisco contents in eucalypt canopies as affected by Acacia neighbourhood. Plant Physiol. Biochem. 34: 753–760.Google Scholar
- Kuppers, M. 1984. Carbon relations and competition between woody species in a central european hedgerow. 3. Carbon and water-balance on the leaf level. Oecologia 65: 94–100.CrossRefGoogle Scholar
- Kuppers, M. 1985. Carbon relations and competition between woody species in a Central European hedgerow. 4. Growth form and partitioning. Oecologia 66: 343–352.CrossRefGoogle Scholar
- Legare, S., Y. Bergeron and D. Pare. 2002. Influence of forest composition on understory cover in boreal mixedwood forests of western Quebec. Silva Fennica 36: 353–366.CrossRefGoogle Scholar
- Li, M.H., G. Hoch and C. Körner. 2001. Spatial variability of mobile carbohydrates within Pinus cembra trees at the alpine treeline. Phyton-Annales Rei Botanicae 41: 203–213.Google Scholar
- Li, M.H., G. Hoch and C. Körner. 2002. Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline. Trees-Structure and Function 16: 331–337.CrossRefGoogle Scholar
- Li, M.H., W.F. Xiao, P. Shi, S.G. Wang, Y.D. Zhong, X.L. Liu, X.D. Wang, X.H. Cai and Z.M. Shi. 2008a. Nitrogen and carbon source-sink relationships in trees at the Himalayan treelines compared to lower elevations. Plant, Cell Environ. 31: 1377–1387.CrossRefGoogle Scholar
- Li, M.H., W.F. Xiao, S.G. Wang, G.W. Cheng, P. Cherubini, X.H. Cai, X.L. Liu, X.D. Wang and W.Z. Zhu. 2008b. Mobile carbohydrates in Himalayan treeline trees I. Evidence for carbon gain limitation but not for growth limitation. Tree Physiol. 28: 1287–1296.CrossRefGoogle Scholar
- Linden, M. and E. Agestam. 2003. Increment and yield in mixed and monoculture stands of Pinus sylvestris and Picea abies based on an experiment in southern Sweden. Scandinavian J. Forest Res. 18: 155–162.CrossRefGoogle Scholar
- Lloret, F., J. Penuelas and M. Estiarte. 2005. Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland. J. Veg. Sci. 16: 67–76.CrossRefGoogle Scholar
- Makinen, H. and A. Isomaki. 2004. Thinning intensity and growth of Scots pine stands in Finland. Forest Ecol. Manage. 201: 311–325.CrossRefGoogle Scholar
- Manceur, A.M., G.J. Boland, N.V. Thevathasan and A.M. Gordon. 2009. Dry matter partitions and specific leaf weight of soybean change with tree competition in an intercropping system. Agro-forestry Syst. 76: 295–301.CrossRefGoogle Scholar
- Martindale, W. and R.C. Leegood. 1997. Acclimation of photosynthesis to low temperature in Spinacia oleracea L. 2. Effects of nitrogen supply. J. Exp. Bot. 48: 1873–1880.CrossRefGoogle Scholar
- Matsushima, M. and S.X. Chang. 2006. Vector analysis of under-story competition, N fertilization, and litter layer removal effects on white spruce growth and nutrition in a 13-year-old plantation. Forest Ecol. Manage. 236: 332–341.CrossRefGoogle Scholar
- Matsushima, M. and S.X. Chang. 2007. Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass. Plant and Soil 292: 243–258.CrossRefGoogle Scholar
- McDowell, N., J.R. Brooks, S.A. Fitzgerald and B.J. Bond. 2003. Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions. Plant Cell Environ. 26: 631–644.CrossRefGoogle Scholar
- McKenzie, D., C.B. Halpern and C.R. Nelson. 2000. Overstory influences on herb and shrub communities in mature forests of western Washington, USA. Can. J. Forest Res. 30: 1655–1666.CrossRefGoogle Scholar
- Medhurst, J.L. and C.L. Beadle. 2005. Photosynthetic capacity and foliar nitrogen distribution in Eucalyptus nitens is altered by high-intensity thinning. Tree Physiol. 25: 981–991.CrossRefGoogle Scholar
- Miller, B.J., P.W. Clinton, G.D. Buchan and A.B. Robson. 1998. Transpiration rates and canopy conductance of Pinus radiata growing with different pasture understories in agroforestry systems. Tree Physiol. 18: 575–582.CrossRefGoogle Scholar
- Misson, L., C. Vincke and F. Devillez. 2003. Frequency responses of radial growth series after different thinning intensities in Norway spruce (Picea abies (L.) Karst.) stands. Forest Ecol. Manage. 177: 51–63.CrossRefGoogle Scholar
- Mohammed, G.H., T.L. Noland and R.G. Wagner. 1998. Physiological perturbation in jack pine (Pinus banksiana Lamb.) in the presence of competing herbaceous vegetation. Forest Ecol. Manage. 103: 77–85.CrossRefGoogle Scholar
- Montgomery, R.A., P.B. Reich and B.J. Palik. 2010. Untangling positive and negative biotic interactions: views from above and below ground in a forest ecosystem. Ecology 91: 3641–3655.CrossRefGoogle Scholar
- Munguia-Rosas, M.A. and V.J. Sosa. 2008. Nurse plants vs. nurse objects: Effects of woody plants and rocky cavities on the recruitment of the Pilosocereus leucocephalus columnar cactus. Ann. Bot. 101: 175–185.CrossRefGoogle Scholar
- Nambiar, E.K.S. and R. Sands. 1993. Competition for water and nutrients in forests. Can. J. Forest Res. 23: 1955–1968.CrossRefGoogle Scholar
- Nambiar, E.K.S. and P.G. Zed. 1980. Influence of weeds on the water potential, nutrient content and growth of young Radiata pine. Australian Forest Res. 10: 279–288.Google Scholar
- Nichols, J.D., M. Bristow and J.K. Vanclay. 2006. Mixed-species plantations: Prospects and challenges. Forest Ecol. Manage. 233: 383–390.CrossRefGoogle Scholar
- O’Brien, M.J., K.L. O’Hara, N. Erbilgin and D.L. Wood. 2007. Overstory and shrub effects on natural regeneration processes in native Pinus radiata stands. Forest Ecol. Manage. 240: 178–185.CrossRefGoogle Scholar
- Oakley, B.B., M.P. North and J.F. Franklin. 2006. Facilitative and competitive effects of a N-fixing shrub on white fir saplings. Forest Ecol. Manage. 233: 100–107.CrossRefGoogle Scholar
- Oren, R., R.H. Waring, S.G. Stafford and J.W. Barrett. 1987. Twenty-four years of Ponderosa pine growth in relation to canopy leaf area and understory competition. Forest Sci. 33: 538–547.Google Scholar
- Parker, W.C. and D.C. Dey. 2008. Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods. Tree Physiol. 28: 797–804.CrossRefGoogle Scholar
- Parrotta, J.A. 1999. Productivity, nutrient cycling, and succession in single- and mixed-species plantations of Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala in Puerto Rico. Forest Ecol. Manage. 124: 45–77.CrossRefGoogle Scholar
- Perry, M.E.L., W.H. Schacht, G.A. Ruark and J.R. Brandle. 2009. Tree canopy effect on grass and grass/legume mixtures in eastern Nebraska. Agroforestry Syst. 77: 23–35.CrossRefGoogle Scholar
- Petersen, T.D., M. Newton and S.M. Zedaker. 1988. Influence of Ceanothus velutinus and associated forbs on the water-stress and stemwood production of Douglas-fir. Forest Sci. 34: 333–343.Google Scholar
- Ram, J., A. Kumar and J. Bhatt. 2004. Plant diversity in six forest types of Uttaranchal, Central Himalaya, India. Current Sci. 86: 975–978.Google Scholar
- Richards, A.E., D.I. Forrester, J. Bauhus and M. Scherer-Lorenzen. 2010. The influence of mixed tree plantations on the nutrition of individual species: a review. Tree Physiol. 30: 1192–1208.CrossRefGoogle Scholar
- Richards, A.E. and S. Schmidt. 2010. Complementary resource use by tree species in a rain forest tree plantation. Ecol. Appl. 20: 1237–1254.CrossRefGoogle Scholar
- Roberts, S.D. and C.A. Harrington. 2008. Individual tree growth response to variable-density thinning in coastal Pacific Northwest forests. Forest Ecol. Manage. 255: 2771–2781.CrossRefGoogle Scholar
- Rothe, A. and D. Binkley. 2001. Nutritional interactions in mixed species forests: a synthesis. Can. J. Forest Res. 31: 1855–1870.CrossRefGoogle Scholar
- Sagar, R., A.S. Raghubanshi and J.S. Singh. 2008. Comparison of community composition and species diversity of understorey and overstorey tree species in a dry tropical forest of northern India. J. Environ. Manage. 88: 1037–1046.CrossRefGoogle Scholar
- Sala, A., G.D. Peters, L.R. McIntyre and M.G. Harrington. 2005. Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season. Tree Physiol. 25: 339–348.CrossRefGoogle Scholar
- Sands, R. and E.K.S. Nambiar. 1984. Water relations of Pinus radiata in competition with weeds. Can. J. Forest Res. 14: 233–237.CrossRefGoogle Scholar
- Shainsky, L.J., M. Newton and S.R. Radosevich. 1992. Effects of intra-specific and inter-specific competition on root and shoot biomass of young douglas-fir and red alder. Can. J. Forest Res. 22: 101–110.CrossRefGoogle Scholar
- Shainsky, L.J. and C.L. Rose. 1995. Effects of competition on the foliar chemistry of young Douglas-fir in monoculture and mixed stands with young red alder. Can. J. Forest Res. 25: 1969–1977.CrossRefGoogle Scholar
- Silvertown, J. 2004. Plant coexistence and the niche. Trends Ecol. Evol. 19: 605–611.CrossRefGoogle Scholar
- Simard, S.W., T. Blenner-Hassett and I.R. Cameron. 2004. Pre-commercial thinning effects on growth, yield and mortality in even-aged paper birch stands in British Columbia. Forest Ecol. Manage. 190: 163–178.CrossRefGoogle Scholar
- Specht, A. and J. Turner. 2006. Foliar nutrient concentrations in mixed-species plantations of subtropical cabinet timber species and their potential as a management tool. Forest Ecol. Manage. 233: 324–337.CrossRefGoogle Scholar
- Suzuki, A. 2002. Influence of shoot architectural position on shoot growth and branching patterns in Cleyera japonica. Tree Physiol. 22: 885–890.CrossRefGoogle Scholar
- Takahashi, K. 1997. Regeneration and coexistence of two subalpine conifer species in relation to dwarf bamboo in the understorey. J. Veg. Sci. 8: 529–536.CrossRefGoogle Scholar
- Takahashi, K., S. Uemura, J.I. Suzuki and T. Hara. 2003. Effects of understory dwarf bamboo on soil water and the growth of over-story trees in a dense secondary Betula ermanii forest, northern Japan. Ecol. Res. 18: 767–774.Google Scholar
- Thelin, G., U. Rosengren, I. Callesen and M. Ingerslev. 2002. The nutrient status of Norway spruce in pure and in mixed-species stands. Forest Ecol. Manage. 160: 115–125.CrossRefGoogle Scholar
- Tian, D.L., Y.Y. Peng, W.D. Yan, X. Fang, W.X. Kang, G.J. Wang and X.Y. Chen. 2010. Effects of thinning and litter fall removal on fine root production and soil organic carbon content in masson pine plantations. Pedosphere 20: 486–493.CrossRefGoogle Scholar
- Tremmel, D.C. and F.A. Bazzaz. 1995. Plant architecture and allocation in different neighborhoods — Implications for competitive success. Ecology 76: 262–271.CrossRefGoogle Scholar
- Tripathi, S.K., A. Sumida, K. Ono, H. Shibata, S. Uemura, K. Takahashi and T. Hara. 2006. The effects of understorey dwarf bamboo (Sasa kurilensis) removal on soil fertility in a Betula ermanii forest of northern Japan. Ecol. Res. 21: 315–320.CrossRefGoogle Scholar
- Tripathi, S.K., A. Sumida, H. Shibata, S. Uemura, K. Ono and T. Hara. 2005. Growth and substrate quality of fine root and soil nitrogen availability in a young Betula ermanii forest of northern Japan: Effects of the removal of understory dwarf bamboo (Sasa kurilensis). Forest Ecol. Manage. 212: 278–290.CrossRefGoogle Scholar
- Umeki, K. 1995. Importance of crown position and morphological plasticity in competitive interaction in a population of Xanthium canadense. Ann. Bot. 75: 259–265.CrossRefGoogle Scholar
- van der Heijden, M.G.A. and T.R. Horton. 2009. Socialism in soil? The importance of mycorrhizal fungal networks for facilitation in natural ecosystems. J. Ecol. 97: 1139–1150.CrossRefGoogle Scholar
- Vandermeer, J. 1989. The Ecology of Intercropping. Cambridge University Press, New York.CrossRefGoogle Scholar
- Velazquezmartinez, A., D.A. Perry and T.E. Bell. 1992. Response of aboveground biomass increment, growth efficiency, and foliar nutrients to thinning fertilization, and pruning in young Douglas-fir plantations in the Central Oregon Cascades. Can. J. Forest Res. 22: 1278–1289.CrossRefGoogle Scholar
- Wang, X.L., J. Zhao, J.P. Wu, H. Chen, Y.B. Lin, L.X. Zhou and S.L. Fu. 2011. Impacts of understory species removal and/or addition on soil respiration in a mixed forest plantation with native species in southern China. Forest Ecol. Manage. 261: 1053–1060.CrossRefGoogle Scholar
- Watt, M.S., D. Whitehead, e.g., Mason, B. Richardson and M.O. Kimberley. 2003. The influence of weed competition for light and water on growth and dry matter partitioning of young Pinus radiata, at a dryland site. Forest Ecol. Manage. 183: 363–376.CrossRefGoogle Scholar
- Weiner, J. and L. Fishman. 1994. Competition and allometry in Kochia scoparia. Ann. Bot. 73: 263–271.CrossRefGoogle Scholar
- Weiskittel, A.R., L.S. Kenefic, R.S. Seymour and L.M. Phillips. 2009. Long-term effects of precommercial thinning on the stem dimensions, form and branch characteristics of red spruce and balsam fir crop trees in Maine, USA. Silva Fennica 43: 397–409.CrossRefGoogle Scholar
- Wilson, J.B. 1990. Mechanisms of species coexistence - 12 explanations for Hutchinson paradox of the plankton — evidence from New-Zealand plant-communities. New Zealand J. Ecol. 13: 17–42.Google Scholar
- Yildiz, O., K. Cromack, S.R. Radosevich, M.A. Martinez-Ghersa and J.E. Baham. 2011. Comparison of 5th-and 14th-year Douglas-fir and understory vegetation responses to selective vegetation removal. Forest Ecol. Manage. 262: 586–597.CrossRefGoogle Scholar
- Youngberg, C.T. 1975. Effects of fertilization and thinning on growth of ponderosa-pine. Soil Sci. Soc. Am. J. 39: 137–139.CrossRefGoogle Scholar
- Zhang, J.Y., G.W. Cheng, F.H. Yu, N. Krauchi and M.H. Li. 2008. Intensity and importance of competition for a grass (Festuca rubra) and a legume (Trifolium pratense) vary with environmental changes. J. Integrative Plant Biol. 50: 1570–1579.CrossRefGoogle Scholar
- Zhang, J.Y., G.W. Cheng, F.H. Yu, N. Krauchi and M.H. Li. 2009a. Interspecific variations in responses of Festuca rubra and Trifolium pratense to a severe clipping under environmental changes. Biologia 64: 292–298.Google Scholar
- Zhang, S.Y., G. Chauret and Q.J. Tong. 2009b. Impact of precom-mercial thinning on tree growth, lumber recovery and lumber quality in Abies balsamea. Scandinavian J. Forest Res. 24: 425–433.CrossRefGoogle Scholar
- Zhao, J., X.L. Wang, Y.H. Shao, G.L. Xu and S.L. Fu. 2011. Effects of vegetation removal on soil properties and decomposer organisms. Soil Biol. Biochem. 43: 954–960.CrossRefGoogle Scholar
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