Community Ecology

, Volume 6, Issue 1, pp 49–56 | Cite as

Leaf anatomical plasticity of Brachypodium pinnatum (L.) Beauv. growing in contrasting microenvironments in a semiarid loess forest-steppe vegetation mosaic

  • A. MojzesEmail author
  • T. Kalapos
  • K. Virágh
Open Access


After clearcutting xerothermic oakwoods once natural in the forest-steppe loess regions of Hungary, the perennial understorey grass Brachypodium pinnatum has been persisting for decades by establishing microhabitats from shade to full sun. In this paper, we explore variation in leaf anatomy for plants growing in different microhabitat light regimes (full shade under oak canopy, half shade near shrubs, and in unshaded grassland) in situ, and for plants reciprocally transplanted between these microhabitats. Leaf lamina thickness and mesophyll thickness were about 1.5 times greater in the grassland in situ than in oak subcanopy due to an additional layer of mesophyll cells and to 25–32% taller mesophyll cells. Mesophyll thickness and the proportion of veins plus sclerenchyma were lower for plants transplanted from either full or half shade to full sun than in situ plants in the grassland. Parenchymatous bundle sheath tended to be thicker in the grassland than in the two other microhabitats. Mean intervenial distance remained invariable among microsites. These adjustments in leaf anatomy may be a considerable part, but presumably not the dominant component of the medium-term (one year) light acclimation of B. pinnatum and the species’ success in microsites with contrasting light climate appearing side-by-side during secondary vegetation succession.


Leaf structure Light acclimation Mesophyll Phenotypic plasticity Sun and shade leaves Transplant experiment 



Electron transport rate


Leaf mass per area


  1. Boardman, N. K. 1977. Comparative photosynthesis of sun and shade plants. Annu. Rev. Plant Physiol. 28:355–377.CrossRefGoogle Scholar
  2. Bobbink, R. 1991. Effects of nutrient enrichment in Dutch chalk grassland. J. Appl. Ecol. 28:28–41.CrossRefGoogle Scholar
  3. Bobbink, R. and J. H. Willems. 1987. Increasing dominance of Brachypodium pinnatum (L.) Beauv. in chalk grasslands: a threat to a species-rich ecosystem. Biol. Conserv. 40:301–314.CrossRefGoogle Scholar
  4. Castro-Díez, P., J. P. Puyravaud and J. H. C. Cornelissen. 2000. Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types. Oecologia 124:476–486.CrossRefGoogle Scholar
  5. Chazdon, R. L. and S. Kaufmann. 1993. Plasticity of leaf anatomy of two rain forest shrubs in relation to photosynthetic light acclimation. Funct. Ecol. 7:385–394.CrossRefGoogle Scholar
  6. Choong, M. F., P. W. Lucas, J. S. Y. Ong, B. Pereira, H. T. W. Tan and I. M. Turner. 1992. Leaf fracture toughness and sclerophylly: their correlations and ecological implications. New Phytol. 121:597–610.CrossRefGoogle Scholar
  7. De Kroon, H. and J. Knops. 1990. Habitat exploration through morphological plasticity in two chalk grassland perennials. Oikos 59:39–49.CrossRefGoogle Scholar
  8. De Kroon, H. and F. Schieving. 1990. Resource partitioning to clonal growth in relation to clonal growth strategy. In: J. Van Groenendael and H. De Kroon (eds.), Clonal Growth in Plants: Regulation and Function. SPB Acad. Publ., The Hague. pp. 79–94.Google Scholar
  9. Fekete, G. and J. Szujkó-Lacza. 1973. Leaf anatomical and photosynthetical reactions of Quercus pubescens Willd. to environmental factors in various ecosystems. I. Leaf anatomical reactions. Acta Bot. Acad. Sci. Hung. 18(1–2):59–89.Google Scholar
  10. Fekete, G., K. Virágh, R. Aszalós and L. Orlóci. 1998. Landscape and coenological differentiation of Brachypodium pinnatum grasslands in Hungary. Coenoses 13(1):39–53.Google Scholar
  11. Fekete G., K. Virágh, R. Aszalós and I. Précsényi. 2000. Static and dynamic approaches to landscape heterogeneity in the Hungarian forest-steppe zone. J. Veg. Sci. 11:375–382.CrossRefGoogle Scholar
  12. Garnier, E. and G. Laurent. 1994. Leaf anatomy, specific mass and water content in congeneric annual and perennial grass species. New Phytol. 128:725–736.CrossRefGoogle Scholar
  13. Givnish, J. T. 1988. Adaptation to sun and shade: a whole-plant perspective. Aust. J. Plant Physiol. 15:63–92.Google Scholar
  14. GraphPad Software. 2000. GraphPad Instat version 3.05 for Windows 95/NT. GraphPad Software, Inc., San Diego, California, USA, Scholar
  15. Hlwatika, C. N. M. and R. B. Bhat. 2002. An ecological interpretation of the difference in leaf anatomy and its plasticity in contrasting tree species in Orange Kloof, Table Mountain, South Africa. Ann. Bot. 89:109–114.CrossRefGoogle Scholar
  16. Hurst, A. and E. John. 1999. The biotic and abiotic changes associated with Brachypodium pinnatum dominance in chalk grassland in South-East England. Biol. Conserv. 88:75–84.CrossRefGoogle Scholar
  17. Klich, M. G. 2000. Leaf variations in Elaeagnus angustifolia related to environmental heterogeneity. Environ. Exp. Bot. 44:171–183.CrossRefGoogle Scholar
  18. Lambers, H., F. S. Chapin III and T. L.Pons. 1998. Plant Physiological Ecology. Springer-Verlag, New York.CrossRefGoogle Scholar
  19. Laboratory Imaging. 1997. Lucia M 3.52a image analysis software. Laboratory Imaging Ltd, Praha, Czech Republic.Google Scholar
  20. Mendes, M. M., L. C. Gazarini and M. L. Rodrigues. 2001. Acclimation of Myrtus communis to contrasting Mediterranean light environments – effects on structure and chemical composition of foliage and plant water relations. Environ. Exp. Bot. 45:165–178.CrossRefGoogle Scholar
  21. Mojzes, A., T. Kalapos and K. Virágh. 2003. Plasticity of leaf and shoot morphology and leaf photochemistry for Brachypodium pinnatum (L.) Beauv. growing in contrasting microenvironments in a semiarid loess forest-steppe vegetation mosaic. Flora 198:304–320.CrossRefGoogle Scholar
  22. Molnár, E., I. Bagi, Zs. Csintalan and A. Nyakas. 2000. The invasion success of a native grassland species in the Great Hungarian Plain. In: D. Ferenciková, N. Gáborcik, L. Ondrasek, E. Uhlrarová and M. Zimková (eds.), Grassland Ecology V. Proceedings of the 5th Ecological Conference, 23–25 November 1999. Grassland and Mountain Agriculture Research Institute, Banská Bystrica. pp. 423–433.Google Scholar
  23. Niinemets, Ü. 1999. Components of leaf dry mass per area – thickness and density – alter leaf photosynthetic capacity in reverse directions in woody plants. New Phytol. 144:35–47.CrossRefGoogle Scholar
  24. Niinemets, Ü. and J. D. Tenhunen. 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. 20:845–866.CrossRefGoogle Scholar
  25. Oguchi, R., K. Hikosaka and T. Hirose. 2003. Does the photosynthetic light-acclimation need change in leaf anatomy? Plant, Cell Environ. 26:505–512.CrossRefGoogle Scholar
  26. Sage, R. F. 2001. Environmental and evolutionary preconditions for the origin and diversification of the C4 photosynthetic syndrome. Plant biol. 3:202–213.CrossRefGoogle Scholar
  27. Sims, D. A. and R. W. Pearcy. 1992. Response of leaf anatomy and photosynthetic capacity in Alocasia macrorrhiza (Araceae) to a transfer from low to high light. Am. J. Bot. 79(4):449–455.CrossRefGoogle Scholar
  28. Utrillas, M. J. and L. Alegre. 1997. Impact of water stress on leaf anatomy and ultrastructure in Cynodon dactylon (L,) Pers, under natural conditions. Int. J. Plant Sci. 158(3):313–324.CrossRefGoogle Scholar
  29. Van Arendonk, J. J. C. M. and H. Poorter. 1994. The chemical composition and anatomical structure of leaves of grass species differing in relative growth rate. Plant, Cell Environ. 17:963–970.CrossRefGoogle Scholar
  30. Virágh, K. and S. Bartha. 1998. Interspecific associations in different successional stages of Brachypodium pinnatum grassland after deforestation in Hungary. Tiscia 31:3–12.Google Scholar
  31. Zólyomi, B. and G. Fekete. 1994. The Pannonian loess steppe: differentiation in space and time. Abstracta Botanica 18(1):29–41.Google Scholar

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© Akadémiai Kiadó, Budapest 2005

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Authors and Affiliations

  1. 1.Department of Plant Taxonomy and EcologyEötvös Loránd UniversityBudapestHungary
  2. 2.Institute of Ecology and BotanyHungarian Academy of SciencesVácrátótHungary

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