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Biomass and Morphology Responses of Fritillaria unibracteata to Shading and Warming

Haixia Guo, Fusun Shi, Cong Wu, Bo Xu, Yan Wu , Ning Wu
Applied Ecology and Environmental Sciences. 2017, 5(1), 19-23. DOI: 10.12691/aees-5-1-3
Published online: May 24, 2017

Abstract

Fritillaria unibracteata Hsiao and Hsia (Liliaceae) (F. unibracteata) is a perennial and protected species distributed in the meadow or under the shrub at the eastern Tibetan Plateau. To understand how F. unibracteata response to changing environment, OTC (open top chamber) and PSN (Polypropylene shading nets) were used to simulate warming and shading effects. Two years later, dry biomass of each organ (leaf, stem, bulb and root), leaf length (LL), leaf area (LA) and special leaf area (SLA) were measured individually to determine F. unibracteata responses to environmental change. The results showed that: (1) total biomass (TB), leaf biomass (LB) and root biomass (RB) increased significantly under warming treatment (P<0.05), but no significant change under shading; (2) LB/RB was decreased under warming, while both LB/RB and AB/BeB showed no significant responses to shading; (3) leaf morphological characteristics responded significantly to warming and shading; leaf length (LL) significantly increased under both shading and warming treatments (P<0.05); leaf area (LA) increased significantly under warming treatment; special leaf area (SLA) increased significantly under shading treatment. Based on the results above, we get conclusions as below: (1) F. unibracteata responded to warming and shading effects differently; (2) For perennial alpine plants such as F. unibracteata, morphological traits may be more sensitive to environment variations than other traits.

1. Introduction

Biomass allocation is one of the most important adaptation mechanisms of plants. Due to different physical separation of resources between above and under-ground, plants may allocate their biomass differently to organs for different functions. From an economical viewpoint, plants would keep a higher root allocation in response to under-ground resource like water and nutrients stress on one hand, while on the other, a higher shoot allocation to increase their capability to acquire limited resources like light. This flexibility in allocation of biomass is important for plants to maximize their growth rate and survive in the environment with variable resource availability 1, 2, 3.

Despite these earlier works, this notion has been questioned. For example, Wright and Mcconnaughay 4 and Štěpán et al. 5 found that variation of allocation among plants was influenced by both resource variation and ontogenetic drift, plant development would muddy or magnify the biomass allocation observed in experiment. In addition, although allocation variation appears to contribute more to the high diversity in ecological strategies than physiological mechanisms at cellular or molecular levels, there are evidences supporting that morphological characteristics such as leaf area ratio (LAR) and special leaf area (SLA) are more sensitive and important in plants’ adjustment to environmental variation 6, 7. Despite these studies, it still remains unclear how some plant species adjust their biomass allocation to varying resource. Although biomass allocation is the best index to characterize plant’s response to environmental variation, it is still not fully understood for various reasons.

Fritillaria unibracteata Hsiao and Hsia (Liliaceae) is one of the traditional medicine plants ‘Chuanbeimu’ functioning at relieving cough and eliminating phlegm 8. It is a perennial herb growing at an attitude of 3200~4500 m a.s.l. in a shady and humid habitat. In recent years, because of over-harvest and impacts of climate change, the population of F. unibracteata has been threatened. According to the investigations by Xu et al. 9, 10, the biomass, biomass allocation and morphological characteristics (eg, leaf area and special leaf area) of F. Unibracteata were influenced by environment factors such as altitude, but changes of morphological characteristics were more consisted with environment variation. In order to understand how F. unibracteata adapt to environment changes, this research was undertaken to scientifically understand the response of this species to alpine environment in terms of its adaptation of biomass and morphology to warming and shading treatments.

2. Materials and Methods

2.1. Experimental Site

This experiment was carried out in the Chuanbeimu Research Station located in Songpan County, Sichuan Province, China (32°56′18" N, 103°42′45″E) where the elevation is 3,300 m a.s.l. Climate at the experimental site is characterized by cold winter and mild summer. Annual mean temperature is 2.8°C, with -7.6°C minimum monthly mean temperature in January, and 11.7°C maximum monthly mean temperature in July. Water is not a limiting factor for plant growth since the annual mean precipitation is as high as 717.7 mm, and 80% of which falls in the form of rain during the growing season from May to October.

2.2. Material and Treatment

The growth of F. unibracteata comprises three main phases (named as phase A, B and C) in line with its morphological development. Phase A (one year old) is marked by needle shaped leaves. Phase B, two or three years old, the plant has only one leaf and no erected stem. Phase C, four or more years old, the plant exhibits erected stem with prominent leaves. In this study, individuals which had grown two years were selected as the subjects.

Plants with similar sizes were carefully transplanted into three experimental fields, each of which had three 1.5×1.5 m2 plots. In every plot, plants were randomly planted at a distance of 10 cm. Three treatments (including the control, warming and shading) were started respectively in three fields immediately after transplantation. Warming was simulated with open top chambers (OTC) made from Polymethy lmethacrylate plates, 1.5×1.5×1.5 m3. Shading treatment was simulated with Polypropylene shade nets. The net was supported at 70 cm height above the ground. The light below the net was 40% of full light.

2.3. Harvest and Measurement

After two-years, five living plants were harvested from each plot. The plants under different treatments were carefully dug out and washed. After that, each plant was divided into four parts: root, bulb, stem and leaf. Two leaf morphological characteristics, i.e. leaf lengths and leaf area, were measured by electronic vernier and leaf scanner (Canon, LiDE 220) respectively. Then all the parts of plant were oven-dried at 80°C for 48 h and weighed.

2.4. Statistical Analysis

The analyses was carried out using “Statistical Package for Social Sciences” program (SPSS, 16.0) at P=0.05 level. One-way ANOVA was used to analyze the differences of biomass, biomass allocation, leaf length (LL), leaf area (LA) and special leaf area (SLA) among three treatments. Here biomass includes the total biomass (TB), root biomass (RB), bulb biomass (BB), stem biomass (SB) and leaf biomass (LB). Biomass allocation refers to the ratio of leaf biomass to root biomass (LB/RB) and above-ground biomass (leaf and stem biomass) to below-ground biomass (root and bulb biomass) (AB/BeB).

3. Results

3.1. Biomass and Its Allocation

Warming significantly increased F. unibracteata root biomass (RB) and leaf biomass (LB), and correspondingly induced a significant increase of total biomass (TB) (P<0.05). On the contrast, shading treatment showed no significant influence on biomass (see Figure 1).

Resources variation not only led to the change of F. unibracteata biomass, but also shifted its allocation. However, in this experiment, shading did not change LM/RM or AM/BM. Moreover, responses of LB/RB and AB/BeB were different under warming treatment. LB/RB decreased significantly, while AB/BeB showed no obvious change, which suggesting the different significance of LB/RB and AB/BeB on F. unibracteata functions.

3.2. Variations of Leaf Morphological Characteristics

Shading and warming treatments dramatically influenced F. unibracteata leaf morphological characteristics. As shown in Figure 3, leaf length (LL) significantly increased under both shading and warming treatments (P<0.05); leaf area (LA) increased significantly under warming treatment, while leaf special area (LSA) increased significantly under shading treatment.

4. Discussion

4.1. Responses of Biomass and Its Allocation to Warming and Shading Treatments

Total biomass increased significantly under warming treatment compared with the controlled treatment, indicating that the growth of F. unibracteata in this area was temperature-limited. Temperature increase in the future may be good for the growth of F. unibracteata. On the other hand, total biomass showed an increase under shading, but the response was not statistically significant. It was suggested that shade-tolerant species were capable of maintaining a uniform pattern of growth and metabolism over a wide range of light intensities 11, 12. Although light is limit under shading treatment, for shade-tolerant species, reduced respiration rate and saturate photosynthesis point could trade off the reduction caused by light limitation or even enhance plant production 13, 14. This was confirmed by studying results from Li et al. 15 that shading was beneficial for the growth of Fritillaria cirrhosa.

Biomass allocation was one of the most important strategies for plant to adapt to environment change. Significant decrease of LB/RB was induced by warming (P<0.05). Effect of temperature on growth was greater than on resource acquisition ability. Thus, under a higher temperature, nutrients and water become limit, and more biomass need to be allocated to root to meet the demand for their growth 16. However, shading did not show significant effect on biomass allocation (P<0.05). Geng et al. 17 proposed that since perennial species allocate more to shoots during their growth and development, under above-ground resource limitation, biomass allocation may show no apparent plasticity. So did F. unibracteata, which showed no significant responses to light.

4.2. Morphological Responses to Environmental Variations

Consisting with previous studies, morphological characteristics of F. unibracteata leaf such as leaf length (LL), leaf area (LA) and special leaf area (SLA) responding dramatically to shading or warming treatments 18, 19, 20. A common mechanism of plants is to increase their light interception and photosynthetic rate under shading by increasing LL and SLA 21, 22. Similarly, plants could also enhance its production under the condition of increased temperature by increasing LL and SLA 23.

Changes of leaf morphology are common for plants’ adaption to environmental variation which may be more important than biomass allocation sometimes 7. Our results suggested that morphological adjustment of F. unibracteata leaf was more important than biomass allocation, since the morphological traits of F. unibracteata leaf changed greatly under both shading and warming treatments, while biomass and biomass allocation showed significantly change only under warming treatment.

Besides biomass allocation and morphology adjustments, plants also respond to varying environment through physiological modifications. It was suggested that for shading-tolerant species, morphological traits may be more important than physiological traits in response to environmental variation [24-26] 24. In our study, NAR (net assimilation rate, i.e., total biomass per unit leaf area and unit time) showed no significant difference among three treatments (P>0.05). This is consistent with some other studies, in which it was suggested that plants’ responses to environmental variation were due to morphological adjustment other than physiological processes 27, 28.

In conclusion, just as some other species, morphology of F. unibracteata was most important in its response to environment compared with biomass allocation and physiological adjustments 26, 28.

5. Conclusion

In this study, total biomass allocation and the ratio of leaf to root biomass (LB/RB) all changed significantly under warming and showed no significant change under shading treatment; leaf length (LL) increased significantly under both shading and warming treatments; leaf area showed significant change under warming treatment, while special leaf area showed significant change under shading treatment. Even though biomass, biomass allocation and leaf morphology all showed responses to environment variations, for F. unibracteata adjustment to environmental variation, morphological traits would be more critical than biomass and its allocation, since leaf morphological traits changed dramatically under both warming and shading treatments. For F. unibracteata, morphological traits such as leaf length (LL), leaf area (LA) and specific leaf area (SLA) may be more import for the indication of its adaptation to environment variation.

Acknowledgments

The authors are very grateful to Dr. Krishna Oli for his valuable inputs comments during the manuscript preparation. This study was supported by the Sichuan education department Program (16ZA0366).

References

[1]  Troughton, A., “Further studies on the relationship between shoot and root systems of grasses”, Grass and Forage Science, 15, 41-47. March, 1960.
In article      View Article
 
[2]  Bloom, A. J., F. S. Chapin, and H. A. Mooney, “Resource Limitation in Plants - an Economic Analogy”, Annual Review of Ecology and Systematics, 16, 363-392. November, 1985.
In article      View Article
 
[3]  Tilman, D., Plant strategies and the dynamics and structure of plant communities. Princeton University Press, Princeton, N.J., 1988.
In article      View Article
 
[4]  Wright, S.D. and Mcconnaughay, K. D. M., “Interpreting phenotypic plasticity: the importance of ontogeny”, Plant Species Biology, 17, 119-131. December, 2002.
In article      View Article
 
[5]  Štěpán, J., Patáčová, E. And Klimešová J., “Effects of fertilization and competition on plant biomass allocation and internal resources: does plantago lanceolata follow the rules of economic theory?”, Folia Geobotanica, 49(1), 49-64. May, 2014.
In article      View Article
 
[6]  Poorter, H., and Remkes, C., “Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate”, Oecologia, 83, 553-559. February, 1990.
In article      View Article  PubMed
 
[7]  Poorter, H., Niklas K. J., Reich P. B., Oleksyn J., Poot P., and Mommer L., “Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control”, New Phytologist, 193, 30-50. January, 2011.
In article      View Article  PubMed
 
[8]  Committee of Flora of China, Chinese Academy of Sciences, Science Press, Peking, 2004.
In article      
 
[9]  Xu, B., Wang, J., Guo, H., Shi, F., and Wu, N., “Morphological adaptation of wild Fritillaria unibracteata to alpine conditions in the eastern Qinghai-Tibet Plateau”, Chin J Appl Envirorn Biol, 20(6), 955-961. May, 2014.
In article      View Article
 
[10]  Xu, B., Wang, J. N., Shi, F. S., Gao, J., and Wu, N., “Adaptation of biomass allocation patterns of wild Fritillaria unibracteata to alpine environment in the eastern Qinghai-Xizang Plateau”., Chinese Journal of Plant Ecology, 37(3), 187-196.January, 2013.
In article      View Article
 
[11]  Loach, K., “Shade tolerance in tree seedlings. II. Growth analysis of plants raised under artificial shade”, New Phytologist, 69, 273-286. April, 1970.
In article      View Article
 
[12]  Piper, F. I., “Patterns of carbon storage in relation to shade tolerance in southern South American species”, American Journal of Botany, 102(9), 1442-1452. September, 2015.
In article      View Article  PubMed
 
[13]  Lehto, T. and Grace, J., “Carbon balance of tropical tree seedlings:a comparison of two Species”, New Phytologist, 127, 455-463. June, 1994.
In article      View Article
 
[14]  Sims, L., Pastor J., Lee T., and Dewey B., “Nitrogen, phosphorus and light effects on growth and allocation of biomass and nutrients in wild rice”, Oecologia, 170, 65-76. March, 2012.
In article      View Article  PubMed
 
[15]  Li, X. Dai, Y. and Chen, S., “Growth and physiological characteristics of Fritillaria cirrhosa in response to high irradiance and shade in age-related growth phases”, Environmental and Experimental Botany, 67, 77-83. November, 2009.
In article      View Article
 
[16]  Larigauderie A., Ellis B. A., Mills J. N., and Kummerow J., “The Effect of Root and Shoot Temperatures on Growth of Ceanothus-Greggii Seedlings”, Annals of Botany, 67, 97-101. February, 1991.
In article      View Article
 
[17]  Geng, Y. P., Pan X. Y., Xu C. Y., Zhang W. J., Li B., and Chen J. K., “Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of Alternanthera philoxeroides”, Ecological Research, 22, 255-260. July, 2007.
In article      View Article
 
[18]  Mitchell, K. J., “Influence of Light and Temperature on the Growth of Ryegrass (Lolium spp.): I. Pattern of Vegetative Development”, Physiologia Plantarum, 6, 21-46. January, 1953.
In article      View Article
 
[19]  Huante, P. and Rincon E., “Responses to light changes in tropical deciduous woody seedlings with contrasting growth rates”, Oecologia, 113, 53-66. December, 1997.
In article      View Article  PubMed
 
[20]  Markesteijn L. and Poorter L., “Seedling root morphology and biomass allocation of 62 tropical tree species in relation to drought- and shading-tolerance”, Journal of Ecology, 97, 311-325. March, 2009.
In article      View Article
 
[21]  Toledo-Aceves T. and Swaine M. D., “Biomass allocation and photosynthetic responses of lianas and pioneer tree seedlings to light”, Acta Oecologica-International Journal of Ecology, 34, 38-49. July-August, 2008.
In article      View Article
 
[22]  Bing Y. and Wang J., “Effect of long-term warming on growth and biomass allocation of Abies faxoniana seedlings”, Acta Ecologica Sinica, 30(21), 5994-6000. January, 2010.
In article      
 
[23]  Wang C. and Wang S., “A review of research on responses of leaf traits to climate Change”, Chinese Journal of Plant Ecology, 39: 206-216. January, 2015.
In article      View Article
 
[24]  Walters M. B. and Reich P. B., “Low-light carbon balance and shading tolerance in the seedlings of woody plants: do winter deciduous and broad-leaved evergreen species differ?”, New Phytologist, 143, 143-154. July, 1999.
In article      View Article
 
[25]  Portsmuth A. and Niinemets U., “Interacting controls by light availability and nutrient supply on biomass allocation and growth of Betula pendula and B. pubescens seedlings”, Forest Ecology and Management, 227, 122-134. May, 2006.
In article      View Article
 
[26]  Allred B. W., Fuhlendorf S. D., Monaco T. A., and Will R. E., “Morphological and physiological traits in the success of the invasive plant Lespedeza cuneata”, Biological Invasions, 12, 739-749. May, 2009.
In article      View Article
 
[27]  Wright I. J. and Westoby M., “Understanding seedling growth relationships through specific leaf area and leaf nitrogen concentration: generalisations across growth forms and growth irradiance”, Oecologia, 127, 21-29. March, 2001.
In article      View Article
 
[28]  Shipley B. “Net assimilation rate, specific leaf area and leaf mass ratio: which is most closely correlated with relative growth rate?”, A meta-analysis, Functional Ecology, 20, 565-574. June, 2006.
In article      View Article
 

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Cite this article:

Normal Style
Haixia Guo, Fusun Shi, Cong Wu, Bo Xu, Yan Wu, Ning Wu. Biomass and Morphology Responses of Fritillaria unibracteata to Shading and Warming. Applied Ecology and Environmental Sciences. Vol. 5, No. 1, 2017, pp 19-23. http://pubs.sciepub.com/aees/5/1/3
MLA Style
Guo, Haixia, et al. "Biomass and Morphology Responses of Fritillaria unibracteata to Shading and Warming." Applied Ecology and Environmental Sciences 5.1 (2017): 19-23.
APA Style
Guo, H. , Shi, F. , Wu, C. , Xu, B. , Wu, Y. , & Wu, N. (2017). Biomass and Morphology Responses of Fritillaria unibracteata to Shading and Warming. Applied Ecology and Environmental Sciences, 5(1), 19-23.
Chicago Style
Guo, Haixia, Fusun Shi, Cong Wu, Bo Xu, Yan Wu, and Ning Wu. "Biomass and Morphology Responses of Fritillaria unibracteata to Shading and Warming." Applied Ecology and Environmental Sciences 5, no. 1 (2017): 19-23.
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  • Figure 1. Biomass variations under three treatments: the control (CK), shading treatment (ST) and warming treatment (WT). Vertical bars represent S.E. (n =15)
  • Figure 2. Biomass allocation under three treatments: the control (CK), shading treatment (ST) and warming treatment (WT). Vertical bars represent S.E. (n =15)
  • Figure 3. Leaf length (a), leaf area (b) and special leaf area (c) variations under three treatments: the control (CK), shading treatment (ST) and warming treatment (WT). Vertical bars represent S.E. (n =15)
  • Figure 4. Net assimilation rate (total biomass per unit leaf area and unit time) under three treatments: the control (CK), shading treatment (ST) and warming treatment (WT). Vertical bars represent S.E. (n =15)
[1]  Troughton, A., “Further studies on the relationship between shoot and root systems of grasses”, Grass and Forage Science, 15, 41-47. March, 1960.
In article      View Article
 
[2]  Bloom, A. J., F. S. Chapin, and H. A. Mooney, “Resource Limitation in Plants - an Economic Analogy”, Annual Review of Ecology and Systematics, 16, 363-392. November, 1985.
In article      View Article
 
[3]  Tilman, D., Plant strategies and the dynamics and structure of plant communities. Princeton University Press, Princeton, N.J., 1988.
In article      View Article
 
[4]  Wright, S.D. and Mcconnaughay, K. D. M., “Interpreting phenotypic plasticity: the importance of ontogeny”, Plant Species Biology, 17, 119-131. December, 2002.
In article      View Article
 
[5]  Štěpán, J., Patáčová, E. And Klimešová J., “Effects of fertilization and competition on plant biomass allocation and internal resources: does plantago lanceolata follow the rules of economic theory?”, Folia Geobotanica, 49(1), 49-64. May, 2014.
In article      View Article
 
[6]  Poorter, H., and Remkes, C., “Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate”, Oecologia, 83, 553-559. February, 1990.
In article      View Article  PubMed
 
[7]  Poorter, H., Niklas K. J., Reich P. B., Oleksyn J., Poot P., and Mommer L., “Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control”, New Phytologist, 193, 30-50. January, 2011.
In article      View Article  PubMed
 
[8]  Committee of Flora of China, Chinese Academy of Sciences, Science Press, Peking, 2004.
In article      
 
[9]  Xu, B., Wang, J., Guo, H., Shi, F., and Wu, N., “Morphological adaptation of wild Fritillaria unibracteata to alpine conditions in the eastern Qinghai-Tibet Plateau”, Chin J Appl Envirorn Biol, 20(6), 955-961. May, 2014.
In article      View Article
 
[10]  Xu, B., Wang, J. N., Shi, F. S., Gao, J., and Wu, N., “Adaptation of biomass allocation patterns of wild Fritillaria unibracteata to alpine environment in the eastern Qinghai-Xizang Plateau”., Chinese Journal of Plant Ecology, 37(3), 187-196.January, 2013.
In article      View Article
 
[11]  Loach, K., “Shade tolerance in tree seedlings. II. Growth analysis of plants raised under artificial shade”, New Phytologist, 69, 273-286. April, 1970.
In article      View Article
 
[12]  Piper, F. I., “Patterns of carbon storage in relation to shade tolerance in southern South American species”, American Journal of Botany, 102(9), 1442-1452. September, 2015.
In article      View Article  PubMed
 
[13]  Lehto, T. and Grace, J., “Carbon balance of tropical tree seedlings:a comparison of two Species”, New Phytologist, 127, 455-463. June, 1994.
In article      View Article
 
[14]  Sims, L., Pastor J., Lee T., and Dewey B., “Nitrogen, phosphorus and light effects on growth and allocation of biomass and nutrients in wild rice”, Oecologia, 170, 65-76. March, 2012.
In article      View Article  PubMed
 
[15]  Li, X. Dai, Y. and Chen, S., “Growth and physiological characteristics of Fritillaria cirrhosa in response to high irradiance and shade in age-related growth phases”, Environmental and Experimental Botany, 67, 77-83. November, 2009.
In article      View Article
 
[16]  Larigauderie A., Ellis B. A., Mills J. N., and Kummerow J., “The Effect of Root and Shoot Temperatures on Growth of Ceanothus-Greggii Seedlings”, Annals of Botany, 67, 97-101. February, 1991.
In article      View Article
 
[17]  Geng, Y. P., Pan X. Y., Xu C. Y., Zhang W. J., Li B., and Chen J. K., “Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of Alternanthera philoxeroides”, Ecological Research, 22, 255-260. July, 2007.
In article      View Article
 
[18]  Mitchell, K. J., “Influence of Light and Temperature on the Growth of Ryegrass (Lolium spp.): I. Pattern of Vegetative Development”, Physiologia Plantarum, 6, 21-46. January, 1953.
In article      View Article
 
[19]  Huante, P. and Rincon E., “Responses to light changes in tropical deciduous woody seedlings with contrasting growth rates”, Oecologia, 113, 53-66. December, 1997.
In article      View Article  PubMed
 
[20]  Markesteijn L. and Poorter L., “Seedling root morphology and biomass allocation of 62 tropical tree species in relation to drought- and shading-tolerance”, Journal of Ecology, 97, 311-325. March, 2009.
In article      View Article
 
[21]  Toledo-Aceves T. and Swaine M. D., “Biomass allocation and photosynthetic responses of lianas and pioneer tree seedlings to light”, Acta Oecologica-International Journal of Ecology, 34, 38-49. July-August, 2008.
In article      View Article
 
[22]  Bing Y. and Wang J., “Effect of long-term warming on growth and biomass allocation of Abies faxoniana seedlings”, Acta Ecologica Sinica, 30(21), 5994-6000. January, 2010.
In article      
 
[23]  Wang C. and Wang S., “A review of research on responses of leaf traits to climate Change”, Chinese Journal of Plant Ecology, 39: 206-216. January, 2015.
In article      View Article
 
[24]  Walters M. B. and Reich P. B., “Low-light carbon balance and shading tolerance in the seedlings of woody plants: do winter deciduous and broad-leaved evergreen species differ?”, New Phytologist, 143, 143-154. July, 1999.
In article      View Article
 
[25]  Portsmuth A. and Niinemets U., “Interacting controls by light availability and nutrient supply on biomass allocation and growth of Betula pendula and B. pubescens seedlings”, Forest Ecology and Management, 227, 122-134. May, 2006.
In article      View Article
 
[26]  Allred B. W., Fuhlendorf S. D., Monaco T. A., and Will R. E., “Morphological and physiological traits in the success of the invasive plant Lespedeza cuneata”, Biological Invasions, 12, 739-749. May, 2009.
In article      View Article
 
[27]  Wright I. J. and Westoby M., “Understanding seedling growth relationships through specific leaf area and leaf nitrogen concentration: generalisations across growth forms and growth irradiance”, Oecologia, 127, 21-29. March, 2001.
In article      View Article
 
[28]  Shipley B. “Net assimilation rate, specific leaf area and leaf mass ratio: which is most closely correlated with relative growth rate?”, A meta-analysis, Functional Ecology, 20, 565-574. June, 2006.
In article      View Article