Effects of Drought on the Competitive Interference of an Early Successional Species (Rubus fruticosus) on Fagus sylvatica L. Seedlings: ¹⁵N Uptake and Partitioning, Responses of Amino Acids and other N Compounds

Abstract

We assessed the role of water availability as a factor regulating the ability of beech seedlings to cope with competitive interference for nitrogen resources by an early successional species (Rubus fruticosus). A glasshouse experiment was performed with two levels of interference (beech with and without R. fruticosus) and three levels of irrigation (high, intermediate, none). ¹⁵N uptake and partitioning of both species, and composition of N pools in leaves, roots and phloem of beech, were determined. Under all irrigation regimes, ¹⁵N uptake by beech seedlings decreased when grown together with R. fruticosus. R. fruticosus had higher ¹⁵N uptake rates than beech, under all water supply levels. When irrigation was reduced, a substantial decrease in ¹⁵N uptake of beech seedlings and a concurrent increase in ¹⁵N uptake by R. fruticosus were observed. Interference by R. fruticosus and low irrigation also affected the ¹⁵N partitioning in beech seedlings and resulted in reduced allocation of ¹⁵N to the roots. The combination of competitive interference and lack of irrigation led to an increase in soluble non-protein N in roots and leaves of beech, due to protein degradation. This response was attributed to an increase in levels of amino acids serving as osmoprotectants under these conditions. The concentration of proline in leaves of beech was negatively correlated to shoot water potential. A competition-induced reduction of total N in leaves of beech under high and intermediate irrigation was found. These results illustrate (1) the advantage of R. fruticosus in terms of N uptake when compared to young beech, particularly under inadequate water supply, and (2) the changes in N composition of beech seedlings in order to cope with reduced soil water and interference by R. fruticosus.

Key words

Fagus sylvatica - competitive interference - drought - ¹⁵N uptake - ¹⁵N partitioning - amino acids - N compounds

References

• 01 Aerts,  R.. (1999);  Interspecific competition in natural plant communities: mechanisms, trade-offs and plant-soil feedbacks.  Journal of Experimental Botany. 50 29-37
• 02 Aussenac,  G.. (2000);  Interactions between forest stands and microclimate: Ecophysiological aspects and consequences for silviculture.  Annals of Forest Science. 57 287-301
• 03 Ballantyne,  J. S., and Chamberlin,  M. E.. (1994) Regulation of cellular amino acid levels. Cellular and molecular physiology of cell volume regulation. Strange, K., ed. Boca Raton, Florida; CRC Press pp. 111-122
• 04 Baur,  G. A.,, Persson,  H.,, Persson,  T.,, Mund,  M.,, Hein,  M.,, Kummetz,  E.,, Matteucci,  G.,, van Oene,  H.,, Scarascia-Mugnozza,  G.,, and Schulze,  E.-D.. (2000) Linking plant nutrition and ecosystem processes. Carbon and nitrogen in European forest ecosystems. Schulze, E.-D., ed. Berlin; Springer pp. 63-98
• 05 Chang,  S. X.,, Preston,  C. M.,, Mc Cullough,  K.,, Wheetman,  G. F.,, and Barker,  J.. (1996);  Effect of understorey competition on distribution and recovery of ¹⁵N applied to a western red cedar - western hemlock clear-cut site.  Canadian Journal of Forest Research. 26 313-321
• 06 Chiatante,  D.,, Di Iorio,  A.,, Maiuro,  L.,, and Scippa,  S. G.. (1999);  Effect of water stress on root meristems in woody and herbaceous plants during the first stage of development.  Plant and Soil. 217 159-172
• 07 Clinton,  P. W., and Mead,  D. J.. (1994 a);  Competition for nitrogen between Pinus radiata and pasture. I. Recovery of ¹⁵N after one growing season.  Canadian Journal of Forest Research. 24 889-896
• 08 Clinton,  P. W., and Mead,  D. J.. (1994 b);  Competition for nitrogen between Pinus radiata and pasture. II. Trends in plant and soil processes.  Canadian Journal of Forest Research. 24 889-896
• 09 Dambrine,  E.,, Martin,  F.,, Carisey,  N.,, and Granier,  A.. (1995);  Xylem sap composition: A tool for investigating mineral uptake in adult spruce.  Plant and Soil. 168 - 169 233-241
• 10 Dertz,  W.. (1996) Buchenwälder im Zielkatalog der Forstwirtschaft. Buchenwälder - ihr Schutz und ihre Nutzung. Bonn; Stiftung Wald in Not pp. 2-8
• 11 Ellenberg,  H.. (1992) Vegetation Mitteleuropas mit den Alpen. Stuttgart; Eugen Ulmer
• 12 Fotelli,  M. N.,, Geßler,  A.,, Peuke,  A. D.,, and Rennenberg,  H.. (2001);  Drought affects the competition between Fagus sylvatica L. seedlings and an early successional species (Rubus fruticosus): growth, water status and δ¹³C composition.  New Phytologist. 151 427-435
• 13 Gaudet,  C. L., and Keddy,  P. A.. (1988);  A comparative approach to predicting competitive ability from plant traits.  Nature. 334 242-243
• 14 Geßler,  A.,, Schneider,  R.,, Hanemann,  U.,, and Rennenberg,  H.. (1998);  Soluble N compounds in trees exposed to high loads of N: a comparison between the roots of Norway spruce (Picea abies [L.] Karst) and beech (Fagus sylvatica L.) trees grown under field conditions.  New Phytologist. 138 385-399
• 15 Geßler,  A.,, Schrempp,  S.,, Matzarakis,  A.,, Meyer,  H.,, Rennenberg,  H.,, and Adams,  M. A.. (2001);  Carbon isotope composition of phloem sap, wood and foliage of beech (Fagus sylvatica L.): effects of water availability and radiation during the growing season.  New Phytologist. 150 653-664
• 16 Girousse,  C.,, Bournoville,  R.,, and Bonnemain,  J.-L.. (1996);  Water deficit induced changes in concentrations in proline and some other amino acids in the phloem sap of alfalfa.  Plant Physiology. 111 109-113
• 17 Handa,  S., Handa,  A. K.,, Hasegawa,  P. M.,, and Bressan,  R. A.. (1986);  Proline accumulation and the adaptation of cultured plant cells to water stress.  Plant Physiology. 80 938-945
• 18 Hayashi,  H., and Chino,  M.. (1985);  Nitrate and other anions in rice phloem sap.  Plant and Cell Physiology. 26 325-330
• 19 Heuer,  B.. (1994) Osmoregulatory role of proline in water- and salt-stressed plants. Handbook of plant and crop stress. Pessarakli, M., ed. New York; Marcel Dekker Inc. pp. 363-381
• 20 Imsande,  J., and Touraine,  B.. (1994);  N demand and the regulation of nitrate uptake.  Plant Physiology. 105 3-7
• 21 IPCC. . (1997) The Regional Impacts of Climate Change: An Assessment of Vulnerability.  A Special Report of IPCC Working Group II.: Cambridge University Press pp. 517
• 22 IPCC. . (2001) Climate Change 2001: Impacts, Adaptation and Vulnerability.  http://www.IPCC.ch/wg2SPM.pdf
• 23 Johnson,  J. D.,, Tognetti,  R.,, Michelozzi,  M.,, Pinzautti,  S.,, Minotta,  G.,, and Borgetti,  M.. (1997);  Ecophysiological responses of Fagus sylvatica L. seedlings to changing light conditions II. The interaction of light environment and soil fertility on seedling physiology.  Physiologia Plantarum. 101 124-134
• 24 Kelly,  J. M.,, Graves,  W. R.,, and Aiello,  A.. (2000);  Nitrate uptake kinetics for rooted cuttings of Acer rubrum L.  Plant and Soil. 221 221-230
• 25 Kosola,  K. R., and Gross,  K. L.. (1999);  Resource competition and suppression of plants colonizing early successional old fields.  Oecologia. 118 69-75
• 26 Kronzucker,  H. J.,, Siddiqi,  M. Y.,, and Glass,  A. D. M.. (1996);  Kinetics of NH4+ influx in spruce.  Plant Physiology. 110 773-779
• 27 Lea,  P. J., and Ireland,  R. J.. (1999) Nitrogen metabolism in higher plants. Plant amino acids: Biochemistry and biotechnology. Singh, B. K., ed. New York; Marcel Dekker Inc. pp. 1-47
• 28 Lichtenthaler,  H. K., and Wellburn,  A. R.. (1983);  Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents.  Biochemical Society Transactions. 603 591-592
• 29 McNeil,  S. D.,, Nuccio,  M. L.,, and Hanson,  A. D.. (1999);  Betaines and related osmoprotectans. Targets for metabolic engineering of stress resistance.  Plant Physiology. 120 945-949
• 30 Ministerium für Ländlichen Raum, Ernährung, Landwirschaft und Forsten in Baden-Württemberg  (Hrsg.). (1997) Richtlinien zur Jungbestandspflege. 
• 31 Mizunaga,  H.. (2000);  Prediction of PPFD variance at forest floor in a thinned Japanese cypress plantation.  Forest Ecology and Management. 126 309-319
• 32 Nernberg,  D., and Dale,  M. R. T.. (1997);  Competition of five native prairie grasses with Bromus inermis under three moisture regimes.  Canadian Journal of Botany. 75 2140-2145
• 33 Plassard,  C.,, Scheromm,  P.,, Mousain,  D.,, and Salsac,  L.. (1991);  Assimilation of mineral nitrogen and ion balance in the two partners of ectomyccorhizal symbiosis: data and hypothesis.  Experimentia. 47 340-349
• 34 Rennenberg,  H.,, Schneider,  S.,, and Weber,  P.. (1996);  Analysis of uptake and allocation of nitrogen and sulphur compounds by trees in the field.  Journal of Experimental Botany. 47 1491-1498
• 35 Rhodes,  D.,, Handa,  S.,, and Bressan,  R. A.. (1986);  Metabolic changes associated with adaptation of plant cells to water stress.  Plant Physiology. 82 890-903
• 36 Ross,  C. W.. (1992) Assimilation of nitrogen and sulfur. Plant physiology. Salisbury, F. B. and Ross, C. W., eds. Belmont, California; Wadsworth Publishing Company pp. 289-307
• 37 Samaras,  Y.,, Bressan,  R.,, Csonka,  L.,, Carcia-Rios,  M. G.,, Paino D'Urzo,  M.,, and Rhodes,  D.. (1995) Proline accumulation during drought and salinity. Environment and plant metabolism: flexibility and acclimation. Smirnoff, N., ed. Oxford; Bios Scientific pp. 161-187
• 38 Santa Regina,  I., and Tarazona,  T.. (2001);  Nutrient cycling in a natural beech forest and adjacent planted pine in northern Spain.  Forestry. 74 11-28
• 39 Schäfer,  H.-E.. (1995) Materials on the situation of biodiversity in Germany.  Bonn, Germany; Landwirschaftverlag GmbH
• 40 Schneider,  A.,, Geßler,  A.,, Weber,  P.,, van Sengbusch,  D.,, Hanemann,  U.,, and Rennenberg,  H.. (1996);  Soluble N compounds in trees exposed to high loads of N: a comparison of spruce (Picea abies) and beech (Fagus sylvatica) grown under field conditions.  New Phytologist. 134 103-114
• 41 Smethurst,  P. J., and Nambiar,  E. K. S.. (1989);  Role of weeds in the management of nitrogen in a young Pinus radiata plantation.  New Forests. 3 203-224
• 42 Staples,  E. T.,, van Rees,  K. C. J.,, and van Kessel,  C.. (1999);  Nitrogen competition using ¹⁵N between early successional plants and planted white spruce seedlings.  Canadian Journal of Forest Research. 29 1282-1289
• 43 Stewart,  G. R., and Larher,  F.. (1980) Accumulation of amino acids and related compounds in relation to environmental stress. The biochemistry of plants, Vol. 5, Amino acids and derivatives . Miflin, B. J., ed. New York; Academic Press pp. 609-635
• 44 Tarp,  P.,, Helles,  F.,, Holten-Andersen,  P.,, Larsen,  J. B.,, and Strange,  N.. (2000);  Modeling near-natural silvicultural regimes for beech - an economic sensitivity analysis.  Forest Ecology and Management. 130 187-198
• 45 Thibodeau,  L.,, Raymond,  P.,, Camire,  C.,, and Munson,  A. D.. (2000);  Impact of precommercial thinning in balsam fir stands on soil nitrogen dynamics, microbial biomass, decomposition, and foliar nutrition.  Canadian Journal of Forest Research.. 30 229-238
• 46 Tognetti,  R.,, Minotta,  G.,, Pinzauti,  S.,, Michelozzi,  M.,, and Borghetti,  M.. (1998);  Acclimation to changing light conditions of long-term shade-grown beech (Fagus sylvatica L.) seedlings of different geographic origins.  Trees. 12 326-333
• 47 Van der Werf,  A.,, van Nuenen,  M.,, Visser,  A. J.,, and Lambers,  H.. (1993);  Contribution of physiological and morphological plant traits to a species' competitive ability at high and low nitrogen supply - A Hypothesis for inherently fast- and slow-growing monocotyledonous supply.  Oecologia. 94 434-440
• 48 Warren,  S. L.,, Skroch,  W. A.,, and Hinesley,  L. E.. (1987);  Effect of root competition and nitrogen on growth and mineral content of Abies fraseri. .  Canadian Journal of Forest Research. 17 1092-1099
• 49 Wellburn,  F. A. M.,, Lau,  K. K.,, Milling,  P. M. K.,, and Wellburn,  A. R.. (1996);  Drought and air pollution affect nitrogen cycling and free radical scavenging in Pinus halepensis (Mill.).  Journal of Experimental Botany. 47 1361-1367
• 50 Woods,  P. V.,, Nambiar,  E. K. S.,, and Smethurst,  P. J.. (1992);  Effect of annual weeds on water and nitrogen availability to Pinus radiata in young plantation.  Forest Ecology and Management. 48 145-163

Section Editor: U. Lüttge