Abstract
Relationship between leaf orientation in natural or semi-natural herbaceous communities and the local abundance of direct light has rarely been investigated. We present evidence from a permanent plot experiment that seasonal direct light availability in 40 × 40 cm plots significantly affects mean leaf orientation in a semi-natural wooded meadow. In sunny years and in well-illuminated conditions, the leaf surface is exposed predominantly to the north, and vice versa. Two groups of species are responsible for such a response: of the variation in leaf orientation, 41 % is explainable through direct light availability for light-demanding species, and 26 % for graminoid species. In shade-tolerant species and in forbs there is no sign of a regression between leaf orientation and direct light abundance. Ability to plastically react to variation in seasonal and local direct light availability appears to be a trait under considerable selective pressure only in light-demanding plants, and more characteristic of species with graminoid growth form.
Key words
Grassland - direct light availability - leaf orientation - herbaceous canopy
References
-
1
Anderson, M. C..
(1966);
Stand structure and light penetration. II. A theoretical analysis.
Journal of Applied Ecology.
3
41-54
-
2
Baldocchi, D. D.,, Hutchinson, B. A.,, Matt, D. R.,, and McMillen, R. T..
(1985);
Canopy radiative transfer models for spherical and known leaf inclination angle distributions: A test in an oak-hickory.
Journal of Applied Ecology.
22
539-555
-
3
Berg, V. S., and Heuchelin, S..
(1990);
Leaf orientation of soybean seedlings. I. Effect of water potential and photosynthetic photon flux density on paraheliotropism.
Crop Science.
30
631-638
-
4
Caldwell, M. M.,, Meister, H.-P.,, Tenhunen, J. D.,, and Lange, O. L..
(1986);
Canopy structure, light microclimate and leaf gas exchange of Quercus coccifera L. in a Portugese macchia: measurements in different canopy layers and simulations with a canopy model.
Trees.
1
25-41
-
5
Campbell, G. S..
(1986);
Extinction coefficients for radiation in plant canopies calculated using an ellipsoidal inclination angle distribution.
Agricultural and Forest Meteorology.
36
317-321
-
6
Eek, L., and Zobel, K..
(1997);
Effects of additional illumination and fertilization on seasonal changes in fine-scale community structure.
Journal of Vegetation Science.
8
255-234
-
7
Eek, L., and Zobel, K. .
(2000);
Structure and diversity of a species-rich grassland community, treated with additional illumination, fertilization and mowing.
Ecography.
24
157-164
-
8 Ehleringer, J. R., and Forseth, I. N.. (1989) Diurnal leaf movements and productivity in canopies. In Plant canopies: their growth, form and function. Russell, G., Marshall, B., and Jarvis, P. G., eds. Cambridge; Cambrigde Univ. Press pp. 129-142
-
9
Ellenberg, H.,, Weber, H. E.,, Düll, R.,, Wirth, V.,, Werner, W.,, Paulissen, D..
(1991);
Ziegerwerte von Pflanzen in Mitteleuropa.
Scripta Geobotanica.
18
1-248
-
10
Gibson, D.,, Casal, J. J.,, and Deregibus, V. A..
(1992);
The effects of plant density on shoot and leaf lamina angles in Lolium multiflorum and Paspalum dilatatum.
Annals of Botany .
70
69-73
-
11
Goudriaan, J..
(1988);
The bare bones of leaf-angle distribution in radiation models for canopy photosynthesis and energy exchange.
Agricultural and Forest Meteorology.
43
155-169
-
12
Herbert, T. J..
(1991);
Variation in interception of the direct solar beam by top canopy layers.
Ecology.
72
17-22
-
13
Hebert, T. J., and Nilson, T..
(1991);
A model of variance of photosynthesis between leaves and maximization of whole plant photosynthesis.
Photosynthetica.
25
597-606
-
14
Hikosaka, K., and Hirose, T., .
(1997);
Leaf angle as a strategy for light competition: optimal and evolutionary stable light extinction coefficient within a leaf canopy.
Écoscience.
4
501-507
-
15
Kao, W. Y., and Tsai, T. T..
(1998);
Tropic leaf movements, photosynthetic gas exchange, leaf δ13 C and cholorophyll a fluorescence of three soybean species in response to water availability.
Plant, Cell and Environment.
21
1055-1062
-
16
Liira, J., and Zobel, K..
(2000);
Vertical structure of a species-rich grassland canopy, treated with additional illumination, fertilization and mowing.
Plant Ecology.
146
185-195
-
17
Liira, J.,, Zobel, K.,, Mägi, R.,, and Molenberghs, G..
(2002);
Vertical structure of herbaceous canopies: the importance of plant growth-form and species-specific traits.
Plant Ecology,.
in press
-
18
Machado, J.-L., and Reich, P. B..
(1999);
Evaluation of several measures of canopy openness as predictors of photosynthetic photon flux density in deeply shaded conifer-dominated forest understory.
Canadian Journal of Forest Research.
29
1438-1444
-
19
McMillen, G. G., and McClendon, J. H..
(1979);
Leaf angle: an adaptive feature of sun and shade leaves.
Botanical Gazette.
140
437-442
-
20
Myers, D. A.,, Jordan, D. N.,, and Vogelmann, T. C..
(1997);
Inclination of sun and shade leaves influences chloroplast light harvesting and utilization.
Physiologia Plantarum.
99
395-404
-
21
Ryel, R. J., and 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
251-260
-
22
Scott, D., and Wells, J. S..
(1969);
Leaf orientation in barley, lupin, and lucerne stands.
New Zealand Journal of Botany.
7
372-388
-
23
Tappeiner, U., and Cernusca, A..
(1989);
Canopy structure and light climate of different alpine plant communities: analysis by means of a model.
Theoretical and Applied Climatology.
40
81-92
-
24
Urbas, P., and Zobel, K..
(2000);
Adaptive and inevitable morphological plasticity of three herbaceous species in a multi-species community: Field experiment with manipulated nutrients and light.
Acta Oecologica.
21
139-147
-
25
Utsugi, H..
(1999);
Angle distribution of foliage in individual Chamaecyparis obtusa canopies and effect of angle on diffuse light penetration.
Trees - Structure and Function.
14
1-9
-
26
Wang, Y. P., and Jarvis, P. G..
(1988);
Mean leaf angles for the ellipsoidal inclination angle distribution.
Agricultural and Forest Meteorology.
43
319-321
K. Zobel
Department of Botany and Ecology
Tartu University
40 Lai St.
51005 Tartu
Estonia
Email: kzobel@ut.ee
Section Editor: R. Aerts