Chinese Journal of Plant Ecology >
Branching and metabolic exponents in seven woody plants
Received date: 2013-11-19
Accepted date: 2014-05-15
Online published: 2014-06-10
Aims The fractal-like network model of plants (the WBE model) considers that the branching exponents 1/a and 1/b determine the metabolic exponent θ in plants. However, the constant 3/4 metabolic exponent does not completely reflect the plant metabolic scaling. Price, Enquist and Savage extended the WBE model by assuming that branching exponents are not constant but covary with each other, and developed the branching traits covariation model of plants (the PES model). In this paper, we study and compare branching exponents and metabolic exponents in seven woody plants based on leaf area and leaf biomass by using the PES model.
Methods To test the PES model, data on leaf area and leaf biomass of seven woody species were used to determine the values of branching and metabolic exponents. Standardized major axis (SMA) regression protocols were used to determine the numerical values of scaling exponents and normalization constants for each species and across the seven species using the software SMATR. Furthermore, test of a common slope across all species and comparisons between the estimated values and theoretical values proposed by the WBE model were also performed by using the SMATR. Specifically, if the value of p exceeds a critical level, then it is considered that there would be no significant differences among the groups compared and that a common slope can be determined; if the value of p is below the critical level, then it would indicate that the estimated value would be significantly different from the theoretical value.
Important findings Significant allometric relationships between leaf area and leaf biomass were verified within and across species. Specifically, leaf area scaled as 0.86-power of leaf biomass across the entire data sets. Therefore, values of branching and metabolic exponents were determined by using leaf biomass and leaf area separately. Values of the branching exponent 1/a and the metabolic exponent θ based on leaf area were statistically indistinguishable among the seven woody species. On the contrary, values of the branching exponents 1/a and 1/b and the metabolic exponent θ based on leaf biomass differed significantly among the seven woody species. Values of the branching exponents and the metabolic exponent estimated from both leaf area and leaf biomass across the entire data set were all statistically indistinguishable from the theoretical values. Furthermore, compared with the value of metabolic exponent based on leaf biomass, the value of metabolic exponent based on leaf area was statistically more comparable to the theoretical value. The effect of allometric relationship between leaf area and leaf biomass on metabolic rate and relative functional traits of plants should be paid more attention in future research.
Key words: allometry; branching exponent; leaf area; PES model; WBE model
MA Yu-Zhu,CHENG Dong-Liang,ZHONG Quan-Lin,JIN Bing-Jie,LIN Jiang-Ming,LU Hong-Dian,GUO Bing-Qiao . Branching and metabolic exponents in seven woody plants[J]. Chinese Journal of Plant Ecology, 2014 , 38(6) : 599 -607 . DOI: 10.3724/SP.J.1258.2014.00055
[1] | Ackerly DD, Knight CA, Weiss SB, Barton K, Starmer KP (2002). Leaf size, specific leaf area and microhabitat distribution of chaparral woody plants: contrasting patterns in species level and community level analyses. Oecologia, 130, 449-457. |
[2] | Bentley LP, Stegen JC, Savage VM, Smith DD, von Allmen EI, Sperry JS, Reich PB, Enquist BJ (2013). An empirical assessment of tree branching networks and implications for plant allometric scaling models. Ecology Letters, 16, 1069-1078. |
[3] | Brown JH, Gillooly JF, Allen AP, Savage VM, West GB (2004). Toward a metabolic theory of ecology. Ecology, 85, 1771-1789. |
[4] | Cheng DL, Li T, Zhong QL, Wang GX (2010). Scaling relationship between tree respiration rates and biomass. Biology Letters, 6, 715-717. |
[5] | Deng JM, Wang GX, Wei XP (2006). The advance of metabolic regulation studies for macroscopical ecology processes. Acta Ecologica Sinica, 26, 3413-3423. (in Chinese with English abstract) |
[5] | [ 邓建明, 王根轩, 魏小平 (2006). 宏观生态过程的代谢调控研究进展. 生态学报, 26, 3413-3423.] |
[6] | Enquist BJ, Allen AP, Brown JH, Gillooly JF, Kerkhoff AJ, Niklas KJ, Price CA, West GB (2007). Biological scaling: Does the exception prove the rule? Nature, 445, E9-E10. |
[7] | Enquist BJ, Brown JH, West GB (1998). Allometric scaling of plant energetics and population density. Nature, 395, 163-165. |
[8] | Enquist BJ, Niklas KJ (2002). Global allocation rules for patterns of biomass partitioning in seed plants. Science, 295, 1517-1520. |
[9] | Enquist BJ, West GB, Charnov EL, Brown JH (1999). Allometric scaling of production and life-history variation in vascular plants. Nature, 401, 907-911. |
[10] | Falster DS, Warton DI, Wright IJ (2003). (S)MATR: Standardised major axis tests and routines. http://www.bio.mq.edu.au/ecology/SMATR. Cited 2013-05-09. |
[11] | Glazier DS (2005). Beyond the ‘3/4-power law’: variation in the intra-and interspecific scaling of metabolic rate in animals. Biological Reviews, 80, 611-662. |
[12] | Glazier DS (2006). The 3/4-power law is not universal: evolution of isometric, ontogenetic metabolic scaling in pelagic animals. BioScience, 56, 325-332. |
[13] | Glazier DS (2010). A unifying explanation for diverse metabolic scaling in animals and plants. Biological Reviews, 85, 111-138. |
[14] | Koontz TL, Petroff A, West GB, Brown JH (2009). Scaling relations for a functionally two-dimensional plant: Chamaesyce setiloba (Euphorbiaceae). American Journal of Botany, 96, 877-884. |
[15] | Li HT, Han XG, Wu JG (2005). Lack of evidence for 3/4 scaling of metabolism in terrestrial plants. Journal of Integrative Plant Biology, 47, 1173-1183. |
[16] | M?kel? A, Valentine HT (2006). Crown ratio influences allometric scaling in trees. Ecology, 87, 2967-2972. |
[17] | Martin JG, Kloeppel BD, Schaefer TL, Kimbler DL, McNulty SG (1998). Aboveground biomass and nitrogen allocation of ten deciduous southern Appalachian tree species. Canadian Journal of Forest Research, 28, 1648-1659. |
[18] | Mori S, Yamaji K, Ishida A, Prokushkin SG, Masyagina OV, Hagihara A, Hoque ATMR, Suwa R, Osawa A, Nishizono T, Ueda T, Kinjo M, Miyagi T, Kajimoto T, Koike T, Matsuura Y, Toma T, Zyryanova OA, Abaimov AP, Awaya Y, Araki MG, Kawasaki T, Chiba Y, Umari M (2010). Mixed-power scaling of whole-plant respiration from seedings to giant trees. Proceedings of the National Academy of Sciences of the United States of America, 107, 1447-1451. |
[19] | Niklas KJ, Cobb ED (2008). Evidence for “diminishing returns” from the scaling of stem diameter and specific leaf area. American Journal of Botany, 95, 549-557. |
[20] | Niklas KJ, Cobb ED, Niinemets ü, Reich PB, Sellin A, Shipley B, Wright IJ (2007). “Diminishing returns” in the scaling of functional leaf traits across and within species groups. Proceedings of the National Academy of Sciences of the United States of America, 104, 8891-8896. |
[21] | Niklas KJ, Enquist BJ (2001). Invariant scaling relationships for interspecific plant biomass production rates and body size. Proceeding of the National Academy of Sciences of the United States of America, 98, 2922-2927. |
[22] | Pan S, Liu C, Zhang WP, Xu SS, Wang N, Li Y, Gao J, Wang Y, Wang GX (2013). The scaling relationships between leaf mass and leaf area of vascular plant species change with altitude. PLoS ONE, 8, e76872. |
[23] | Peng YH, Niklas KJ, Reich PB, Sun SC (2010). Ontogenetic shift in the scaling of dark respiration with whole-plant mass in seven shrub species. Functional Ecology, 24, 502-512. |
[24] | Price CA, Enquist BJ, Savage VM (2007). A general model for allometric covariation in botanical form and function. Proceedings of the National Academy of Sciences of the United States of America, 104, 13204-13209. |
[25] | Price CA, Ogle K, White EP, Weitz JS (2009). Evaluating scaling models in biology using hierarchical Bayesian approaches. Ecology Letters, 12, 641-651. |
[26] | Reich PB, Tjoelker MG, Machado JL, Oleksyn J (2006). Universal scaling of respiratory metabolism, size and nitrogen in plants. Nature, 439, 457-461. |
[27] | Savage VM, Deeds EJ, Fontana W (2008). Sizing up allometric scaling theory. PLoS Computational Biology, 4, e1000171. |
[28] | Wang JF, Feng YL (2004). The effect of light intensity on biomass allocation, leaf morphology and relative growth rate of two invasive plants. Acta Phytoecologica Sinica, 28, 781-786. (in Chinese with English abstract) |
[28] | [ 王俊峰, 冯玉龙 (2004). 光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响. 植物生态学报, 28, 781-786.] |
[29] | Warton DI, Wright IJ, Falster DS, Westoby M (2006). Bivariate line-fitting methods for allometry. Biological Reviews, 81, 259-291. |
[30] | West GB, Brown JH, Enquist BJ (1997). A general model for the origin of allometric scaling laws in biology. Science, 276, 122-126. |
[31] | West GB, Brown JH, Enquist BJ (1999a). A general model for the structure and allometry of plant vascular systems. Nature, 400, 664-667. |
[32] | West GB, Brown JH, Enquist BJ (1999b). The fourth dimension of life: fractal geometry and allometric scaling of organisms. Science, 284, 1677-1679. |
[33] | Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets ü, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004). The worldwide leaf economics spectrum. Nature, 428, 821-827. |
/
〈 |
|
〉 |