Back
 OJE  Vol.3 No.2 , May 2013
Seasonal patterns of light availability and light use of broadleaf evergreens in a deciduous forest understory: Potential mechanisms for expansion
Abstract: In recent years, expansion of native and exotic evergreen shrubs into forest understories has been documented worldwide. Dense shrub thickets may interfere with tree establishment, suppress herbaceous cover, and contribute substantially to total standing crop of leaf biomass. Expansion may occur because evergreen shrubs exploit seasonal variations in irradiance and temperature that are characteristic of temperate understory environments. We quantified leaf-level light environment and photosynthetic activity of three sympatric broadleaf evergreens (Ilex opaca, Kalmia latifolia, and Myrica cerifera) in a deciduous forest understory in Charles City County,Virginia,USAin order to understand seasonal intra- and interspecific ranges of broadleaf evergreen physiology. Two species (K. latifolia and M. cerifera) represent a diverse taxonomic range within broadleaf evergreens, and often form expansive thickets. We measured parameters related to canopy structure (e.g., bifurcation ratio, leaf angle) and photosynthetic performance (e.g., electron transport rate or ETR, chlorophyll content), to identify potential mechanisms facilitating expansion. ETR varied both seasonally and among species. In summer, M. cerifera ETR was nearly double that ofI. opaca or K. latifolia. Additionally, leaf temperature enhanced photosynthetic capacity of expansive species. Evergreen species, though capable of fixing carbon throughout the year, often exhibit slow growth rates and low physiological activity. Yet, we observed that the range of evergreen physiological activity may be broader than previously recognized. Furthermore, our results indicate potential for changes in composition and expansion of the evergreen shrub layer by species that exhibit structural and physiological mechanisms advantageous for future rises in temperature.
Cite this paper: Shiflett, S. , Zinnert, J. and Young, D. (2013) Seasonal patterns of light availability and light use of broadleaf evergreens in a deciduous forest understory: Potential mechanisms for expansion. Open Journal of Ecology, 3, 151-160. doi: 10.4236/oje.2013.32018.
References

[1]   Vitousek, P.M. and Walker, L.R. (1989) Biological invasion by Myrica faya in Hawai’i: Plant demography, nitrogen fixation, and ecosystem effects. Ecological Monographs, 59, 247-265. doi:10.2307/1942601

[2]   Brothers, T.S. and Spingarn, A. (1992) Forest fragmentation and alien plant invasion of central Indiana old-growth forests. Conservation Biology, 6, 91-100. doi:10.1046/j.1523-1739.1992.610091.x

[3]   Lavergne, C., Rameau, J.-C. and Figier, J. (1999) The invasive woody weed Ligustrum robustum subsp. walkeri threatens native forests on La Réunion. Biological Invasions, 1, 377-392. doi:10.1023/A:1010001529227

[4]   Banasiak, S.E. and Meiners, S.J. (2009) Long term dynamics of Rosa multiflora in a successional system. Biological Invasions, 11, 215-224. doi:10.1007/s10530-008-9226-1

[5]   Mitchell, J.D., Lockaby, B.G. and Brantley, E.F. (2011) Influence of Chinese privet (Ligustrum sinense) on decomposition and nutrient availability in riparian forests. Invasive Plant Science and Management, 4, 437-447. doi:10.1614/IPSM-D-11-00020.1

[6]   Monk, C.D., McGinty, D.T. and Day Jr., F.P. (1985) The ecological importance of Kalmia latifolia and Rhododendron maximum in the deciduous forest of the southern Appalachians. Bulletin of the Torrey Botanical Club, 112, 187-193. doi:10.2307/29-96415

[7]   Beier, C.M., Horton, J.L., Walker, J.F., Clinton, B.D. and Nilsen, E.T. (2005) Carbon limitation leads to suppression of first year oak seedlings beneath evergreen understory shrubs in southern Appalachian hardwood forests. Plant Ecology, 176, 131-142. doi:10.1007/s11258-004-0119-9.

[8]   Dobbs, M.M. and Parker, A.J. (2004) Evergreen understory dynamics in Coweeta forest, North Carolina. Physical Geography, 6, 481-498. doi:10.2747/0272-3646.25.6.481

[9]   Monk, C.D. and Day, F.P. (1985) Vegetation analysis, primary production and selected nutrient budgets for a southern Appalachian oak forest: A synthesis of IBP studies at Coweeta. Forest Ecology and Management, 10, 87-113. doi:10.1016/0378-1127(85)90015-5

[10]   Young, D.R., Shao, G. and Porter, J.H. (1995) Spatial and temporal growth dynamics of barrier island shrub thickets. American Journal of Botany, 82, 628-645. doi:10.2307/2445422

[11]   Kurten, E.L., Snyder, C.P., Iwata, T. and Vitousek, P.M. (2008) Morella cerifera invasion and nitrogen cycling on a lowland Hawaiian lava flow. Biological Invasions, 10, 19-24. doi:10.1007/s10530-007-9101-5

[12]   Zinnert, J.C., Shiflett, S.A., Vick, J.K. and Young, D.R. (2011) Woody vegetative cover dynamics in response to recent climate change on an Atlantic coast barrier island: A remote sensing approach. Geocarto International, 26, 595-612. doi:10.1080/10106049.2011.621031

[13]   Chapin, F.S. III (1980) The mineral nutrition of wild plants. Annual Review of Ecology and Systematics, 11, 233-260. doi:10.1146/annurev.es.11.110180.001313

[14]   Chabot, B.F. and Hicks, D.J. (1982) The ecology of leaf life spans. Annual Review of Ecology and Systematics, 13, 229-259. doi:10.1146/annurev.es.13.110182.001305

[15]   Aerts, R. (1995) The advantages of being evergreen. Trends in Ecology and Evolution, 10, 402-407. doi:10.1016/S0169-5347(00)89156-9

[16]   Baldocchi, D.D., Ma, S., Rambal, S., Misson, L., Ourcival, J.-M., Limousin, J.-M., Pereira, J. and Papale, D. (2010) On the differential advantages of evergreenness and deciduousness in Mediterranean oak woodlands: A flux perspective. Ecological Applications, 20, 1583-1597. doi:10.1890/08-2047.1

[17]   Marty, C., Lamaze, T. and Pornon, A. (2010) Leaf life span optimizes annual biomass production rather than plant photosynthetic capacity in an evergreen shrub. New Phytologist, 87, 407-416. doi:10.1111/j.1469-8137.2010.03290.x

[18]   Letts, M.G., Rodríguez-Calcerrada, J., Rolo, V. and Rambal, S. (2012) Long-term physiological and morphological acclimation by the evergreen shrub Buxus simpervirens L. to understory and canopy gap light intensities. Trees, 26, 479-491. doi:10.1007/s00468-011-0609-z

[19]   Monk, C.D. (1966) An ecological significance of ever-greenness. Ecology, 47, 504-505. doi:10.2307/1932995

[20]   Hollinger, D.Y. (1992) Leaf and simulated whole-canopy photosynthesis in two co-occurring tree species. Ecology, 73, 1-14. doi:10.2307/1938716

[21]   Givnish, T.J. (2002) Adaptive significance of evergreen vs. deciduous leaves: Solving the triple paradox. Silva Fennica, 36, 703-743.

[22]   Muller, O., Hikosaka, K. and Hirose, T. (2005) Seasonal changes in light and temperature affect the balance between light harvesting and light utilization components of photosynthesis in an evergreen understory shrub. Oecologia, 143, 501-508. doi:10.1007/s00442-005-0024-5

[23]   Wang, X., Arora, R., Horner, H.T. and Krebs, S.L. (2008) Structural adaptations in overwintering leaves of thermonastic and nonthermonastic Rhododendron species. Journal of the American Society for Horticultural Science, 133, 768-776.

[24]   Muller, O., Hirose, T., Werger, M.J.A. and Hikosaka, K. (2011) Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub. Annals of Botany, 108, 529-536. doi:10.1093/aob/mcr167

[25]   Chazdon, R.L. (1986) Light variation and carbon gain in rain forest understorey palms. Journal of Ecology, 74, 995-1012. doi:10.2307/2260229

[26]   Chazdon, R.L. and Pearcy, R.W. (1991) The importance of sunflecks for forest understory plants. Bioscience, 41, 760-766. doi:10.2307/1311725

[27]   Neufeld, H.S. and Young, D.R. (2003) Ecophysiology of the herbaceous layer in temperate deciduous forest. In: Gilliam, F.S. and Roberts, M.R., Eds., The Herbaceous Layer in Forest in Eastern North America, Oxford University Press, Oxford, 38-90.

[28]   Roberts, S.W., Knoerr, K.R. and Strain, B.R. (1979) Comparative field water relations of four co-occurring forest tree species. Canadian Journal of Botany, 57, 1876-1882. doi:10.1139/b79-237

[29]   Young, D.R. (1985) Crown architecture, light interception, and stomatal conductance patterns for sympatric deciduous and evergreen species in a forest understory. Canadian Journal of Botany, 63, 2425-2429. doi:10.1139/b85-346

[30]   Krinard, R.M. (1973) American holly: An American wood. United States Department of Agriculture Forest Service. FS-242.

[31]   Valladares, F., Arrieta, S., Aranda, I., Lorenzo, D., Sánchez-Gómez, D., Tena, D., Suárez, F. and Pardos, J.A. (2005) Shade tolerance, photoinhibition sensitivity and phenotypic plasticity of Ilex aquilifolium in continental Mediterranean sites. Tree Physiology, 25, 1041-1052. doi:10.1093/treephys/25.8.1041

[32]   Reich, P.B., Walters, M.B., Ellsworth, D.S., Vose, J.M., Volin, J.C., Gresham, C. and Bowman, W.D. (1998) Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span: A test across biomes and functional groups. Oecologia, 114, 471-482. doi:10.1007/s004420050471

[33]   Hughes, N.M. and Smith, W.K. (2007) Seasonal photosynthesis and anthocyanin production in ten broadleaf evergreen species. Functional Plant Biology, 34, 1072-1079. doi:10.1071/FP07205

[34]   Young, D.R. (1992) Photosynthetic characteristics and potential moisture stress for the actinorhizal shrub, Myrica cerifera (Myricaceae), on a Virginia barrier island. American Journal of Botany, 79, 2-7. doi:10.2307/2445189

[35]   Brantley, S.T. and Young, D.R. (2010) Linking light attenuation, sunflecks and canopy architecture in mesic shrub thicket. Plant Ecology, 206, 225-326. doi:10.1007/s11258-009-9637-9

[36]   Lichtenthaler, H.K. (1987) Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods in Enzymology, 148,350-382. doi:10.1016/0076-6879(87)48036-1

[37]   Schreiber, U., Bilger, W. and Neubauer, C. (1994) Chlorophyll fluorescence as a non-intrusive indicator for rapid assessment of in vivo photosynthesis. In: Schulze, E.D. and Caldwell, M.M., Eds., Ecophysiology of Photosynthesis, Ecological studies, Springer, Berlin, 49-70.

[38]   Jones, T.J., Luton, C.D., Santiago, L.S. and Goldstein, G. (2010) Hydraulic constraints on photosynthesis in subtropical evergreen broad leaf forest and pine woodland trees of the Florida Everglades. Trees, 24, 471-478. doi:10.1007/s00468-010-0415-z

[39]   Schreiber, U., Neubauer, C. and Klughammer, C. (1988) New ways of assessing photosynthetic activity with a pulse modulation fluorometer. In: Lichtenthaler, H.K., Ed., Applications of Chlorophyll Fluorescence, Kluwer Academic Publishing, Dordrecht, 63-69.

[40]   Lovelock, C.E., Osmond, C.B. and Jebb, M. (1994) Photoinhibition and recovery in tropical plant species: Response to disturbance. Oecologia, 97, 297-307.

[41]   Genty, B., Briantais, J.M. and Baker, N.R. (1989) The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochimica et Biophysica Acta, 990, 87-92. doi:10.1016/S0304-4165(89)80016-9

[42]   Demmig-Adams, B., Maguas, C., Adams, W.W. III, Meyer, A., Kilian, E. and Lange, O.L. (1990) Effect of high light on the efficiency of photochemical energy conversion in a variety of lichen species with green and bluegreen phycobionts. Planta, 180, 400-409. doi:10.1007/BF01160396

[43]   Andrews, J.R., Fryer, M.J. and Baker, N.R. (1995) Characterization of chilling effects on photosynthetic performance of maize crops during early season growth using chlorophyll fluorescence. Journal of Experimental Botany, 46, 1195-1203. doi:10.1093/jxb/46.9.1195

[44]   Stemke, J.A. and Santiago, L.S. (2011) Consequences of light absorptance in calculating electron transport rate of desert and succulent plants. Photosynthetica, 49, 195-200. doi:10.1007/s11099-011-0026-y

[45]   Zinnert, J.C., Via, S.M. and Young, D.R. (2013) Distinguishing natural from anthropogenic stress in plants: Physiology, fluorescence and hyperspectral reflectance. Plant and Soil, 366, 133-141. doi:10.1007/s11104-012-1414-1

[46]   Naumann, J.C., Young, D.R. and Anderson, J.E. (2007) Linking leaf chlorophyll fluorescence properties to physiological responses for stress detection in coastal plant species. Physiologia Plantarum, 131, 422-433. doi:10.1111/j.1399-3054.2007.00973.x

[47]   Flores-Moya, A., Hanelt, D., Figueroa, F.-L., Altamirano, M., Vinegla, B. and Salles, S. (1999) Involvement of solar UV-B radiation in recovery of inhibited photosynthesis in the brown alga Dictyota dichotoma (Hudson) Lamouroux. Journal of Photochemistry and Photobiology B: Biology, 49, 129-135. doi:10.1016/S1011-1344(99)00046-9

[48]   Campbell, G.S. (1990) Derivation of an angle density function for canopies with ellipsoidal leaf angle distributions. Agricultural and Forest Meteorology, 49, 173-176. doi:10.1016/0168-1923(90)90030-A

[49]   Sands, P.J. (1995) Modeling canopy production 1: Optimal distribution of photosynthetic resources. Australian Journal of Plant Physiology, 22, 593-601. doi:10.1071/PP9950593

[50]   Drouet, J.L. and Moulia, B. (1997) Spatial re-orientation of maize leaves affected by initial plant orientation and density. Agricultural and Forest Meteorology, 88, 85-100. doi:10.1016/S0168-1923(97)00047-6

[51]   Falster, D.S. and 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

[52]   Teh, C.B.S., Simmonds, L.P. and Wheeler, T.R. (2000) An equation for irregular distributions of leaf azimuth density. Agricultural and Forest Meteorology, 102, 223-234. doi:10.1016/S0168-1923(00)00132-5

[53]   Whitney, G.G. (1976) The bifurcation ratio as an indicator of adaptive strategy in woody plant species. Bulletin of the Torrey Botanical Club, 103, 67-72. doi:10.2307/2484833

[54]   Kempf, J.S. and Pickett, S.T.A. (1981) The role of branch length and angle in branching pattern of forest shrubs along a successional gradient. New Phytologist, 88, 111-116. doi:10.1111/j.1469-8137.1981.tb04574.x

[55]   Brantley, S.T. and Young, D.R. (2009) Contribution of sunflecks is minimal in expanding shrub thickets compared to temperate forest. Ecology, 90, 1021-1029. doi:10.1890/08-0725.1

[56]   Steingraeber, D.A., Kascht, L.J. and Franck, D.H. (1979) Variation in shoot morphology and bifurcation ratio in sugar maple (Acer saccharum) saplings. American Journal of Botany, 66, 441-445. doi:10.2307/2442397

[57]   Demmig-Adams, B., Gilmore, A.M., and Adams, W.W. III. (1996) In vivo functions of carotenoids in higher plants. The FASEB Journal, 10, 403-412.

[58]   Boardman, N.K. (1977) Comparative photosynthesis of sun and shade plants. Annual Review of Plant Physiology, 28, 355-377.doi:10.1146/annurev.pp.28.060177.002035

[59]   Martin, C.E. and Warner, D.A. (1984) The effects of desiccation on concentrations and a/b ratios of chlorophyll in Leucobryum glaucum and Thuidium delicatulum. New Phytologist, 96, 545-550. doi:10.1111/j.1469-8137.1984.tb03588.x

[60]   Dale, M.P. and Causton, D.R. (1992) Use of chlorophyll a/b ratio as a bioassay for the light environment of a plant. Functional Ecology, 6, 190-196. doi:10.2307/2389754

[61]   Walther, G.R. (2000) Climatic forcing on the dispersal of exotic species. Phytocoenologia, 30, 409-430.

[62]   Potter, D.A. and Kimmerer, T.W. (1986) Seasonal allocation of defense investment in Ilex opaca Aiton and constraints on a specialist leafminer. Oecologia, 69, 217-224. doi:10.1007/BF00377625

[63]   Young, D.R. (2007) Estimating aboveground net primary production in shrub-dominated ecosystems. In: Fahey, T.J. and Knapp, A.K., Eds., Principles and standards for measuring primary production, Oxford University Press, NewYork, 49-68. doi:10.1093/acprof:oso/9780195168662.003.0004

[64]   Vick, J.K. (2011) Woody encroachment mechanisms of a symbiotic N-fixing shrub: Ecophysiology, facilitation, and resource use efficiency. PhD dissertation, Virginia Commonwealth University, Richmond.

 
 
Top