Interactions Between Shoot Age Structure, Nutrient Availability and Physiological Integration in the Giant Bamboo Phyllostachys pubescens

Abstract

The age structure of adult shoots, the nutrient availability of the habitat, and their interaction, are important factors influencing the productivity of bamboo groves. In a field fertilization experiment over two years we examined the impact of physiological integration on the emergence, growth, and survival of new shoots of Phyllostachys pubescens, a giant woody bamboo. Impacts of physiological integration were compared using uniform and patchy fertilization treatments.

The number of new shoots emerging per plot significantly increased with the application of fertilizer (NPK), but the increase was constrained by the age structure of adult shoots. In a year with a high proportion of shoots with new (first-year) leaves many more new shoots emerged than in a year with a low proportion of new leaves. Mean survivorship of the new shoots was constant at 20 % for all treatments in both years of study. Surprisingly, fertilization did not increase DBH or height of the new shoots, suggesting that shoot size was fixed early in ontogeny.

The pattern of fertilizer application, either uniform or patchy, did not affect the total number of new shoots produced. In the case of patchy application, more new shoots developed in unfertilized patches in comparison to uniformly unfertilized plots, probably because these parts of the clone received resources via the rhizomes from the adult shoots in adjacent fertilized patches. The production of new shoots in fertilized patches, in turn, was lower than that in uniformly fertilized plots as a result of this physiological integration. The results are discussed in the general context of the impact of integration on clonal plant performance in dense stands.

Key words

Age structure - bamboo - clonal growth - patchy fertilization - physiological integration - vegetative reproduction

References

• 01 Alpert,  P., and Stuefer,  J. F.. (1997) Division of labour in clonal plants. The ecology and evolution of clonal plants. de Kroon, H. and van Groenendael, J., eds. Leiden; Backhuys Publishers pp. 137-154
• 02 Bamboo Research Institute of Nanjing Forestry University. (1974) Cultivation of bamboo groves. Beijing; Agriculture Press p. 278
• 03 Birch,  C. P. D., and Hutchings,  M. J.. (1994);  Exploitation of patchily distributed soil resources by the clonal herb Glechoma hederacea. .  J. Ecol.. 82 653-664
• 04 de Kroon,  H.. (1993);  Competition between shoots in stands of clonal plants.  Plant Species Biol.. 8 85-94
• 05 de Kroon,  H., and Kalliola,  R.. (1995);  Shoot dynamics of the giant grass Gynerium sagittatum in Peruvian Amazon floodplains, a clonal plant that does show self-thinning.  Oecologia. 101 124-131
• 06 Dong,  M.. (1995);  Morphological responses to local light conditions in clonal herbs from contrasting habitats, and their modification due to physiological integration.  Oecologia. 101 282-288
• 07 Fliervoet,  L. M.,, Zhong,  Z. C.,, Liu,  Y. C.,, Miao,  S. L.,, Dong,  M.,, and Werger,  M. J. A.. (1989);  Diversity and above-ground structure of the understorey of Phyllostachys pubescens groves on Mount Jinyun, Sichuan, China.  Flora. 182 203-219
• 08 Geber,  M. A.,, de Kroon,  H.,, and Watson,  M. A.. (1997 a);  Organ preformation in mayapple as a mechanism for historical effects on demography.  J. Ecol.. 85 211-223
• 09 Geber,  M. A.,, Watson,  M. A.,, and de Kroon,  H.. (1997 b) Organ preformation, development, and resource allocation in perennials. Plant resource allocation. Bazzaz, F. A. and Grace, J., eds. New York; Academic Press pp. 113-141
• 10 Hirose,  T., and Werger,  M. J. A.. (1987);  Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation pattern in the canopy.  Oecologia. 72 520-526
• 11 Hsiung,  W. Y.,, Xiao,  J. H.,, and Lou,  Y. P.. (1990);  Development of rhizome-culm systems in a newly established stand of Phyllostachys praecox. .  Bamboo J. (Japan). 8 64-68
• 12 Huang,  Q. M., and Yang,  D. D.. (1989);  Research on photosynthesis of bamboo.  Bamboo Res. (China). 8 (2) 8-18
• 13 Hutchings,  M. J.. (1979);  Weight-density relationships in ramet populations of clonal perennial herbs, with special reference to the - 3/2 power law.  J. Ecol.. 67 21-33
• 14 Hutchings,  M. J., and Price,  E. A. C.. (1993);  Does physiological integration enable clonal plants to integrate the effects of environmental heterogeneity?.  Plant Species Biol.. 8 95-105
• 15 Hutchings,  M. J., and Wijesinghe,  D. K.. (1997);  Patchy habitats, division of labour and growth dividends in clonal plants.  Trends Ecol. Evol.. 12 390-394
• 16 Isagi,  Y.,, Kawahara,  T.,, Kamo,  K.,, and Ito,  H.. (1997);  Net production and carbon cycling in a bamboo Phyllostachys pubescens stand.  Plant Ecol.. 130 41-52
• 17 Janzen,  D. H.. (1976);  Why bamboos wait so long to flower.  Annu. Rev. Ecol. Syst.. 7 347-391
• 18 Jónsdóttir,  I. S., and Watson,  M. A.. (1997) Extensive physiological integration: An adaptive trait in resource-poor environments?. The ecology and evolution of clonal plants. de Kroon, H. and van Groenendael, J., eds. Leiden; Backhuys Publishers pp. 109-136
• 19 Li,  R.,, During,  H. J.,, Werger,  M. J. A.,, and Zhong,  Z. C.. (1998 a);  Positioning of new shoots relative to adult shoots in groves of giant bamboo Phyllostachys pubescens. .  Flora. 193 315-321
• 20 Li,  R.,, Werger,  M. J. A.,, During,  H. J.,, and Zhong,  Z. C.. (1998 b);  Biennial variation in production of new shoots in groves of the giant bamboo Phyllostachys pubescens in Sichuan, China.  Plant Ecol.. 135 103-112
• 21 Li,  R.,, Werger,  M. J. A.,, During,  H. J.,, and Zhong,  Z. C.. (1998 c);  Biomass distribution in a grove of the giant bamboo Phyllostachys pubescens in Chongqing, China.  Flora. 194 89-96
• 22 Li,  R.,, Werger,  M. J. A.,, During,  H. J.,, and Zhong,  Z. C.. (1998 d);  Carbon and nutrient dynamics in relation to growth rhythm in the giant bamboo Phyllostachys pubescens. .  Plant and Soil. 201 113-123
• 23 Liao,  G. L.. (1990);  On the flowering and regeneration of Phyllostachys pubescens. .  Bamboo J. (Japan). 8 21-23
• 24 Liu,  Y. C.,, Fliervoet,  L. M.,, Zhong,  Z. C.,, and Werger,  M. J. A.. (1988) Stand structure of giant bamboo (Phyllostachys pubescens) groves in Sichuan, China. Vegetation structure in relation to carbon and nutrient economy: production, decomposition and atmospheric interception. Verhoeven, J. T. A., Heil, G. W., and Werger, M. J. A., eds. The Hague; SPB Academic Publishing pp. 37-43
• 25 MacClure,  F. A.. (1966) The bamboos: a fresh perspective. Cambridge, Massachusetts; Harvard University Press p. 347
• 26 Marshall,  C.. (1990) Source-sink relations of interconnected ramets. Clonal growth in plants: regulation and function. van Groenendael, J. and de Kroon, H., eds. The Hague; SPB Academic Publishing pp. 23-41
• 27 Mooney,  H. A.. (1986) Photosynthesis. Plant ecology. Crawley, M.J., ed. Oxford; Blackwell Scientific Publications pp. 345-374
• 28 Numata,  M.. (1979) Structure and succession of bamboo vegetation. Ecology of grasslands and bamboolands in the world. Numata, M., ed. The Hague; Dr. W. Junk Publishers pp. 237-258
• 29 Pitelka,  L. F., and Ashmun,  J. W.. (1985) Physiology and integration of ramets in clonal plants. Population biology and evolution of clonal organisms. Jackson, J. B. C., Buss, L. W., and Cook, R. E., eds. New Haven; Yale University Press pp. 399-437
• 30 Shi,  Q. T., and Bian,  Y. R.. (1987);  Study on the application of chemical fertilizers in bamboo stump.  J. Bamboo Res. (in Chinese). 6 24-34
• 31 Sokal,  R. R., and Rohlf,  F. J.. (1995) Biometry, 3d ed. New York; W. H. Freeman and Company p. 887
• 32 StatSoft Inc.. (1993) Statistica for Windows. Tulsa, Oklahoma, USA; Release 4.5
• 33 Stuefer,  J. F.,, de Kroon,  H.,, and During,  H. J.. (1996);  Exploitation of environmental heterogeneity by spatial division of labour in a clonal plant.  Funct. Ecol.. 10 328-334
• 34 Stuefer,  J. F.,, During,  H. J.,, and de Kroon,  H.. (1994);  High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environments.  J. Ecol.. 82 511-518
• 35 Suzuki,  J., and Hutchings,  M. J.. (1997) Interactions between shoots in clonal plants and the effects of stored resources on the structure of shoot populations. The ecology and evolution of clonal plants. de Kroon, H. and van Groenendael, J., eds. Leiden; Backhuys Publishers pp. 311-329
• 36 Ueda,  K.. (1960);  Studies on the physiology of bamboo, with reference to practical application.  Bulletin of the Kyoto University of Forestry. 30 1-169
• 37 Watson,  M. A.. (1984);  Developmental constraints: effect on population growth and patterns of resource allocation in a clonal plant.  Am. Nat.. 123 411-426
• 38 Watson,  M. A.,, Geber,  M. A.,, and Jones,  C. S.. (1995);  Ontogenetic contingency and the expression of plant plasticity.  Trends Ecol. Evol.. 10 474-475
• 39 Wu,  B. S.. (1984);  A preliminary analysis of the relationship between the underground structure of a Phyllostachys pubescens forest and its yield.  J. Bamboo Res. (in Chinese). 3 49-58
• 40 Zhou,  J. Y.,, Hu,  C. Z.,, and Yan,  L. P.. (1985);  Investigations on the rhizomes of bamboo groves for food consumption.  J. Bamboo Res. (in Chinese). 4 57-64

Section Editor: R. Aerts