Response of clonal plasticity of to different canopy conditions of subalpine coniferous forest

Abstract

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments.

References

1. Abe M, Izaki J, Miguchi H, Masaki T, Makita A, Nakashizuka T (2002). The effects of Sasa and canopy gap formation on tree regeneration in an old beech forest. J Veg Sci, 13: 565–574. doi:10.1658/1100-9233(2002)013[0565:TEOSAC]2.0.CO;2
2. Abe M, Miguchi H, Honda A, Makita A, Nakashizuka T (2005). Short-termchanges affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis. J Veg Sci, 16: 49–56. doi:10.1658/1100-9233(2005)016[0049:SCAROF]2.0.CO;2
3. Chen J S, Dong M, Yu D, Liu Q (2004). Clonal architecture and ramet population characteristics of Lysimachia congestiflora growing under differentlight conditions. Chinese Journal of AppliedEcology, 15(8): 1383–1388 (in Chinese)
4. Denslow J S, Newell E, Ellison A M (1991). The effect of understory palms andcyclanths on the growth and survival of Inga seedlings. Biotropica, 23: 225–234. doi:10.2307/2388199
5. Dong M (1996). Clonal growth in plants in relation to resource heterogeneity:foraging behavior. Acta Botanica Sinica, 38(10): 828–835 (in Chinese)
6. Franklin J F, Maeda T, Oshumi T, Matsui M, Yagi H, Hawk G M (1979). Subalpine coniferous forests of central Honshu, Japan. Ecol Monogr, 49: 311–334. doi:10.2307/1942487
7. Griscom B W, Ashton P M S (2003). Bamboocontrol of forest succession: Guadua sarcocarpa in southeastern Peru. For Ecol Manage, 175: 445–454. doi:10.1016/S0378-1127(02)00214-1
8. Holz C A, Veblen T T (2006). Treeregeneration responses to Chusquea montana bamboo die-off in a subalpine Nothofagus forest in the southern Andes. J Veg Sci, 17: 19–28. doi:10.1658/1100-9233(2006)017[0019:TRRTCM]2.0.CO;2
9. Iida S (2004). Indirect negative influence of dwarf bamboo on survivalof Quercus acorn by hoarding behavior of wood mice. For Ecol Manage, 202: 257–263. doi:10.1016/j.foreco.2004.07.022
10. Li Z X, Zheng H (2004). The spatialdistribution characteristics of throughfall under Abies faxoniana forest in the Wolong Nature Reserve. Acta Ecologica Sinica, 24: 1015–1021 (in Chinese)
11. Liu Q, Li Y X, Zhong Z C (2004). Effects of moisture availabilityon clonal growth in bamboo Pleioblastus maculata. Plant Ecol, 173: 107–113. doi:10.1023/B:VEGE.0000026334.40661.06
12. Ma W L, Zhong Z C (1998). Morphologicaladaptability of clonal herb Iris japonica to changed light condition. Chinese Journalof Applied Ecology, 9(1): 23–26 (in Chinese)
13. Nakashizuka T (1987). Regeneration dynamics of beech forests in Japan. Vegetation, 69: 169–175. doi:10.1007/BF00038698
14. Narukawa Y, Yamamoto S (2001). Effectsof dwarf bamboo (Sasa sp.) andforest floor microsites on conifer seedling recruitment in a subalpineforest, Japan.Forest Ecology and Management, 163: 61–70. doi:10.1016/S0378-1127(01)00527-8
15. Noguchi M, Yoshida T (2004). Treeregeneration in partially cut conifer–hardwood mixed forestsin northern Japan: roles of establishment substrate and dwarf bamboo. For Ecol Manage, 190: 335–344. doi:10.1016/j.foreco.2003.10.024
16. Noguchi M, Yoshida T (2005). Factorsinfluencing the distribution of two co-occurring dwarf bamboo species(Sasa kurilensis and S. senanensis) in a conifer-broadleaved mixed stand in northern Hokkaido.Ecol Res, 20: 25–30. doi:10.1007/s11284-004-0009-6
17. Qin Z S, Taylor A H, Cai X S (1993). Bamboo and Forest Dynamic Successionin the Ecological Environment of Giant Panda in Wolong. Beijing: China Forestry Press,13, 30–57, 245–318, 343–375 (in Chinese)
18. Reid D G, Taylor A H, Hu J C, Qin Z S (1991). Environmental influences on Bashania fangiana bamboo growth and implications for giant panda conservation. Journal of Applied Ecology, 28: 855–868 (in Chinese). doi: 10.2307/2404212
19. Song L X, Tao J P, Wang W, Xi Y, Wang Y J, Ran C Y (2006). The ramet population structures of the clonal bamboo Fargesia nitida in different canopy conditionsof subalpine dark coniferous forest in the Wolong Nature Reserve. Acta Ecologica Sinica, 26(3): 730–736 (in Chinese)
20. Tabarelli M, Mantovani W (2000). Gap-phaseregeneration in a tropical montane forest: the effects of gap structureand bamboo species. Plant Ecol, 148(2): 149–155. doi:10.1023/A:1009823510688
21. Takahashi K, Uemura S, Suzuki J I, Hara T (2003). Effects of understory dwarf bamboo on soil water and the growth ofoverstory trees in a dense secondary Betulaermanii forest, northern Japan. Ecol Res, 18: 767–774. doi:10.1007/s11284-003-0594-9
22. Taylor A H, Huang J Y, Zhou S Q (2004). Canopy tree development and undergrowthbamboo dynamics in old-growth Abies–Betula forests in southwestern China: a 12-year study. For Ecol Manage, 200: 347–360. doi:10.1016/j.foreco.2004.07.007
23. Taylor A H, Jang S W, Zhao L J, Liang C P, Miao C J, Huang J Y (2006). Regeneration patterns and tree species coexistence inold-growth Abies–Picea forestsin southwestern China. For Ecol Manage, 223: 303–317. doi:10.1016/j.foreco.2005.11.010
24. Taylor A H, Qin Z (1988a). Regenerationpatterns in old-growth Abies-Betula forests in the Wolong Natural Reserve, Sichuan China. J Ecol, 76: 1204–1218. doi:10.2307/2260643
25. Taylor A H, Qin Z (1988b). Treereplacement patterns in subalpine Abies–Betula forests in Wolong Natural Reserve, China. Vegetation, 78: 141–149. doi:10.1007/BF00033423
26. Taylor A H, Qin Z (1992). Treeregeneration after bamboo die-off in Abies-Betula Forests, Wolong Natural Reserve, China. J Veg Sci, 3: 253–260. doi:10.2307/3235687
27. Taylor A H, Qin Z S, Liu J (1995). Tree regeneration in an Abies faxoniana forest after bamboo dieback,Wang Long Natural Reserve, China. Can JFor Res, 25: 2034–2039. doi:10.1139/x95-220
28. Veblen T T, Veblen A T, Schlegel F M (1979). Understorey patterns in mixed evergreen-deciduous Nothofagus forests in Chile. J Ecol, 67: 809–823. doi:10.2307/2259216
29. Wang J X,Ma Z G (1993). EcologicalStudies on Giant Panda'S Main Food Bamboos. Chengdu: Shichuan Scientific andTechnical Press, 25–29, 62–87 (in Chinese)
30. Wang W, Tao J P, Li Z F, Zhang W Y, Ding Y (2004). Gap features of subalpinedark coniferous forest in Wolong Nature Reserve. Chinese Journal of Applied Ecology, 15(11): 1989–1993 (in Chinese)
31. Widmer Y (1998).Pattern and performance understory bamboos (Chusquea spp.) under different canopy bamboosclosures in old-growth oak forest in Costa Rica.Biotropica, 30(3): 400–415. doi:10.1111/j.1744-7429.1998.tb00074.x
32. Yu X H, Tao J P, Li Y, Wang Y J, Xi Y, Zhang W Y, Zang R G (2006). Ramet populationstructure of Fargesia nitida (Mitford)Keng f. et Yi in different successional stands of the subalpine coniferousforest in Wolong Nature Reserve.J Integr Plant Biol, 48: 1147-1153. doi:10.1111/j.1744-7909.2006.00330.x
33. Zhang S M, Chen Y F, Yu F H, Xing X R, Li L H, Dong M (2003). Clonal growth and clonal morphology of Potentilla reptans var. sericophylla in forest understorey and gap.Acta PhytoecologicaSinica, 27(4): 567–571 (in Chinese)