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Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE

Sexual differences in morphology and aboveground biomass allocation in relation to branch number in Morus alba saplings

Huihui Huan A , Bixia Wang A , Gang Liu B , Xiao Xu A C D and Xinhua He A D
+ Author Affiliations
- Author Affiliations

A Key Laboratory of Southwest China Wildlife Resources Conservation, China West Normal University, Ministry of Education, Nanchong, Sichuan 637 009, China.

B Sericultural Research Institute, Sichuan Academy of Agricultural Sciences, Nanchong, Sichuan 637 000, China.

C College of Life Sciences, China West Normal University, Nanchong, Sichuan 637 009, China.

D Corresponding authors. Email: xuxiao_cwnu@163.com; xinhua.he@uwa.edu.au

Australian Journal of Botany 64(3) 269-275 https://doi.org/10.1071/BT15189
Submitted: 17 August 2015  Accepted: 19 April 2016   Published: 9 May 2016

Abstract

The effect of branch number on crown size and biomass accumulation has been reported for many trees, but knowledge about sexual differences in growth and aboveground biomass allocation related to branch number is still limited. Morus alba L. is a dioecious plant of economic importance, and was employed as a model species in our study. Thirty male and 30 female 1-month-old M. alba saplings with single or multiple (five) primary branches were cultivated for one growing season. Sexual differences in gas exchange, morphology, biomass accumulation and allocation, and relationships among morphological and biomass traits were investigated. Branch multiplication significantly increased the total leaf number, total leaf area, total leaf mass, aboveground mass, leaf : stem biomass ratio and specific leaf area at the plant level but decreased net photosynthetic rate (Pn), leaf number, mean single-leaf area, stem length, basal diameter and branch biomass at the branch level. Five-branch male saplings exhibited a significantly higher Pn, total leaf number, total leaf area, total leaf mass and aboveground biomass than did female ones. No such between plant sex differences were found in single-branch saplings. Further, branch number was positively correlated with aboveground mass in male saplings only. Our results demonstrate that branch multiplication might result in sexual differences in morphology and biomass accumulation and male saplings may allocate greater resources than females to aboveground parts for vegetative growth.

Additional keywords: branch number, dioecious, growth strategy, net photosynthetic rate.


References

Ågren J (1988) Sexual differences in biomass and nutrient allocation in the dioecious Rubus chamaemorus. Ecology 69, 962–973.
Sexual differences in biomass and nutrient allocation in the dioecious Rubus chamaemorus.Crossref | GoogleScholarGoogle Scholar |

Antos JA, Allen GA (1994) Biomass allocation among reproductive structures in the dioecious shrub Oemleria cerasiformis – a functional interpretation. Journal of Ecology 82, 21–29.
Biomass allocation among reproductive structures in the dioecious shrub Oemleria cerasiformis – a functional interpretation.Crossref | GoogleScholarGoogle Scholar |

Barathi P, Sundar D, Reddy AR (2001) Changes in mulberry leaf metabolism in response to water stress. Biologia Plantarum 44, 83–87.
Changes in mulberry leaf metabolism in response to water stress.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3MXjtFygsrc%3D&md5=77ab982c8a3f069998ddef7c6d91f9e3CAS |

Bartelink HH (1997) Allometric relationships for biomass and leaf area of beech (Fagus sylvatica). Annals of Forest Science 54, 39–50.
Allometric relationships for biomass and leaf area of beech (Fagus sylvatica).Crossref | GoogleScholarGoogle Scholar |

Bullock SH (1984) Biomass and nutrient allocation in a neotropical dioecious palm. Oecologia 63, 426–428.
Biomass and nutrient allocation in a neotropical dioecious palm.Crossref | GoogleScholarGoogle Scholar |

Burgess KS, Husband BC (2006) Habitat differentiation and the ecological costs of hybridization: the effects of introduced mulberry (Morus alba) on a native congener (M. rubra). Journal of Ecology 94, 1061–1069.
Habitat differentiation and the ecological costs of hybridization: the effects of introduced mulberry (Morus alba) on a native congener (M. rubra).Crossref | GoogleScholarGoogle Scholar |

Burk TE, Nelson ND, Isebrands JG (1983) Crown architecture of short-rotation, intensively cultured Populus. III. A model of first-order branch architecture. Canadian Journal of Forest Research 13, 1107–1116.
Crown architecture of short-rotation, intensively cultured Populus. III. A model of first-order branch architecture.Crossref | GoogleScholarGoogle Scholar |

Cepeda-Cornejo V, Dirzo R (2010) Sex-related differences in reproductive allocation, growth, defense and herbivory in three dioecious neotropical palms. PLoS One 5, e9824
Sex-related differences in reproductive allocation, growth, defense and herbivory in three dioecious neotropical palms.Crossref | GoogleScholarGoogle Scholar | 20352113PubMed |

Ceulemans R, Hinckley TM, Heilman PE, Isebrands JG, Stettler RF (1989) Crown architecture in relation to productivity of Populus clones in the Pacific Northwest, USA. Annals of Forest Science 46, 199–201.
Crown architecture in relation to productivity of Populus clones in the Pacific Northwest, USA.Crossref | GoogleScholarGoogle Scholar |

Ceulemans R, Stettler RF, Hinckley TM, Isebrands JG, Heilman PE (1990) Crown architecture of Populus clones as determined by branch orientation and branch characteristics. Tree Physiology 7, 157–167.
Crown architecture of Populus clones as determined by branch orientation and branch characteristics.Crossref | GoogleScholarGoogle Scholar | 14972913PubMed |

Chaitanya K, Sundar D, Reddy AR (2001) Mulberry leaf metabolism under high temperature stress. Biologia Plantarum 44, 379–384.
Mulberry leaf metabolism under high temperature stress.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3MXmtlentrs%3D&md5=13c53e1c15ebd2936c69544421e30fe1CAS |

Chaitanya K, Jutur P, Sundar D, Reddy AR (2003) Water stress effects on photosynthesis in different mulberry cultivars. Plant Growth Regulation 40, 75–80.
Water stress effects on photosynthesis in different mulberry cultivars.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXis1CisLw%3D&md5=d204440ccbe8034c298592f365c12a3dCAS |

Chen F, Chen L, Zhao H, Korpelainen H, Li C (2010) Sex-specific responses and tolerances of Populus cathayana to salinity. Physiologia Plantarum 140, 163–173.
Sex-specific responses and tolerances of Populus cathayana to salinity.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXht1ejurnJ&md5=9c49e0997d4ca527604d285e324905b4CAS | 20561244PubMed |

Darwin C (1877) ‘The different forms of flowers on plants of the same species.’ (John Murray: London)

Dawson TE, Ehleringer JR (1993) Gender-specific physiology, carbon isotope discrimination, and habitat distribution in boxelder, Acer negundo. Ecology 74, 798–815.
Gender-specific physiology, carbon isotope discrimination, and habitat distribution in boxelder, Acer negundo.Crossref | GoogleScholarGoogle Scholar |

Delph LF, Knapczyk FN, Taylor DR (2002) Among – population variation and correlations in sexually dimorphic traits of Silene latifolia. Journal of Evolutionary Biology 15, 1011–1020.
Among – population variation and correlations in sexually dimorphic traits of Silene latifolia.Crossref | GoogleScholarGoogle Scholar |

Ercisli S, Orhan E (2007) Chemical composition of white (Morus alba), red (Morus rubra) and black (Morus nigra) mulberry fruits. Food Chemistry 103, 1380–1384.
Chemical composition of white (Morus alba), red (Morus rubra) and black (Morus nigra) mulberry fruits.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXjs1Cqt7g%3D&md5=aef24614c52683fd61af32854a80a32fCAS |

Fisher JB, Honda H (1979) Branch geometry and effective leaf area: a study of terminalia-branching pattern. 1. Theoretical trees. American Journal of Botany 66, 633–644.
Branch geometry and effective leaf area: a study of terminalia-branching pattern. 1. Theoretical trees.Crossref | GoogleScholarGoogle Scholar |

Gehring JL, Linhart YB (1993) Sexual dimorphisms and response to low resources in the dioecious plant Silene latifolia (Caryophyllaceae). International Journal of Plant Sciences 154, 152–162.
Sexual dimorphisms and response to low resources in the dioecious plant Silene latifolia (Caryophyllaceae).Crossref | GoogleScholarGoogle Scholar |

Gholz HL (1982) Environmental limits on aboveground net primary production, leaf area, and biomass in vegetation zones of the Pacific Northwest. Ecology 63, 469–481.
Environmental limits on aboveground net primary production, leaf area, and biomass in vegetation zones of the Pacific Northwest.Crossref | GoogleScholarGoogle Scholar |

Grant MC, Mitton JB (1979) Elevational gradients in adult sex ratios and sexual differentiation in vegetative growth rates of Populus tremuloides Michx. Evolution 33, 914–918.
Elevational gradients in adult sex ratios and sexual differentiation in vegetative growth rates of Populus tremuloides Michx.Crossref | GoogleScholarGoogle Scholar |

Gross KL, Soule JD (1981) Differences in biomass allocation to reproductive and vegetative structures of male and female plants of a dioecious, perennial herb, Silene alba (Miller) Krause. American Journal of Botany 68, 801–807.
Differences in biomass allocation to reproductive and vegetative structures of male and female plants of a dioecious, perennial herb, Silene alba (Miller) Krause.Crossref | GoogleScholarGoogle Scholar |

Harmer R (1992) Relationships between shoot length, bud number and branch production in Quercus petraea (Matt.) Liebl. Forestry 65, 61–72.
Relationships between shoot length, bud number and branch production in Quercus petraea (Matt.) Liebl.Crossref | GoogleScholarGoogle Scholar |

Hoffmann AJ, Alliende MC (1984) Interactions in the patterns of vegetative growth and reproduction in woody dioecious plants. Oecologia 61, 109–114.
Interactions in the patterns of vegetative growth and reproduction in woody dioecious plants.Crossref | GoogleScholarGoogle Scholar |

Huan H, Xu X, Liu G, He J, Yang Y, Huang K (2014) Effect of branch number on the growth and development of Morus alba saplings. Acta Ecologica Sinica 34, 823–831. [in Chinese with English abstract]

Isebrands JG, Nelson ND (1982) Crown architecture of short-rotation, intensively cultured Populus II. Branch morphology and distribution of leaves within the crown of Populus ‘Tristis’ as related to biomass production. Canadian Journal of Forest Research 12, 853–864.
Crown architecture of short-rotation, intensively cultured Populus II. Branch morphology and distribution of leaves within the crown of Populus ‘Tristis’ as related to biomass production.Crossref | GoogleScholarGoogle Scholar |

Kauppi A, Kiviniitty M, Ferm A (1988) Growth habits and crown architecture of Betula pubescens Ehrh. of seed and sprout origin. Canadian Journal of Forest Research 18, 1603–1613.
Growth habits and crown architecture of Betula pubescens Ehrh. of seed and sprout origin.Crossref | GoogleScholarGoogle Scholar |

Kellomäki S (1986) A model for the relationship between branch number and biomass in Pinus sylvestris crowns and the effect of crown shape and stand density on branch and stem biomass. Scandinavian Journal of Forest Research 1, 455–472.
A model for the relationship between branch number and biomass in Pinus sylvestris crowns and the effect of crown shape and stand density on branch and stem biomass.Crossref | GoogleScholarGoogle Scholar |

Kim JH, Kim TJ, Lee CH, Paek KY, Hahn EJ, Kim HH (2003) Effect of number of branches on quality and yield of secondary cut flower production of Dendranthema grandiflorum ‘Herman de Boon’. Korean Journal of Horticultural Science and Technology 21, 212–215.

King DA (1998) Relationship between crown architecture and branch orientation in rain forest trees. Annals of Botany 82, 1–7.
Relationship between crown architecture and branch orientation in rain forest trees.Crossref | GoogleScholarGoogle Scholar |

Kohorn LU (1995) Geographic variation in the occurrence and extent of sexual dimorphism in a dioecious shrub, Simmondsia chinensis. Oikos 74, 137–145.
Geographic variation in the occurrence and extent of sexual dimorphism in a dioecious shrub, Simmondsia chinensis.Crossref | GoogleScholarGoogle Scholar |

Korpelainen H (1992) Patterns of resource allocation in male and female plants of Rumex acetosa and Rumex acetosella. Oecologia 89, 133–139.
Patterns of resource allocation in male and female plants of Rumex acetosa and Rumex acetosella.Crossref | GoogleScholarGoogle Scholar |

Kumar SG, Reddy AM, Sudhakar C (2003) NaCl effects on proline metabolism in two high yielding genotypes of mulberry (Morus alba L.) with contrasting salt tolerance. Plant Science 165, 1245–1251.
NaCl effects on proline metabolism in two high yielding genotypes of mulberry (Morus alba L.) with contrasting salt tolerance.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXos1KntLc%3D&md5=731a7024f13106ad171a222d7749703cCAS |

Lebon E, Pellegrino A, Louarn G, Lecoeur J (2006) Branch development controls leaf area dynamics in grapevine (Vitis vinifera) growing in drying soil. Annals of Botany 98, 175–185.
Branch development controls leaf area dynamics in grapevine (Vitis vinifera) growing in drying soil.Crossref | GoogleScholarGoogle Scholar | 16679414PubMed |

Li C, Ren J, Luo J, Lu R (2004) Sex-specific physiological and growth responses to water stress in Hippophae rhamnoides L. populations. Acta Physiologiae Plantarum 26, 123–129.
Sex-specific physiological and growth responses to water stress in Hippophae rhamnoides L. populations.Crossref | GoogleScholarGoogle Scholar |

Li C, Xu G, Zang R, Korpelainen H, Berninger F (2007) Sex-related differences in leaf morphological and physiological responses in Hippophae rhamnoides along an altitudinal gradient. Tree Physiology 27, 399–406.
Sex-related differences in leaf morphological and physiological responses in Hippophae rhamnoides along an altitudinal gradient.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXkslSmt78%3D&md5=1e54e120750c2984fabe7099141bf6dfCAS | 17241981PubMed |

Luo P, Zhou SL (2007) Effect of land use on ecological benefit of farm belt in suburbs: A case study was conducted of Gaoping District, Nanchong City, Sichuan Province. Journal of Ecology and Rural Environment 23, 6–10. [in Chinese with English abstract]

McDowell SCL, McDowell NG, Marshall JD, Hultine K (2000) Carbon and nitrogen allocation to male and female reproduction in Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca, Pinaceae). American Journal of Botany 87, 539–546.
Carbon and nitrogen allocation to male and female reproduction in Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca, Pinaceae).Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3Mngs1KnsA%3D%3D&md5=0a29ad1bcc26b9f0e7fef0e432d3999cCAS |

Mizumachi E, Mori A, Osawa N, Akiyama R, Tokuchi N (2006) Shoot development and extension of Quercus serrata saplings in response to insect damage and nutrient conditions. Annals of Botany 98, 219–226.
Shoot development and extension of Quercus serrata saplings in response to insect damage and nutrient conditions.Crossref | GoogleScholarGoogle Scholar | 16709576PubMed |

Nakamura Y, Morita A (2006) Effects of pruning of tea cuttings in paper pots on the number of branches and plant growth before and after transplanting. Nihon Sakumotsu Gakkai Kiji 75, 289–295.
Effects of pruning of tea cuttings in paper pots on the number of branches and plant growth before and after transplanting.Crossref | GoogleScholarGoogle Scholar |

Nelson ND, Burk T, Isebrands JG (1981) Crown architecture of short-rotation, intensively cultured Populus. I. Effects of clone and spacing on first-order branch characteristics. Canadian Journal of Forest Research 11, 73–81.
Crown architecture of short-rotation, intensively cultured Populus. I. Effects of clone and spacing on first-order branch characteristics.Crossref | GoogleScholarGoogle Scholar |

Nelson AS, Weiskittel AR, Wagner RG (2014) Development of branch, crown, and vertical distribution leaf area models for contrasting hardwood species in Maine, USA. Trees 28, 17–30.
Development of branch, crown, and vertical distribution leaf area models for contrasting hardwood species in Maine, USA.Crossref | GoogleScholarGoogle Scholar |

Obeso JR, Alvarez-Santullano M, Retuerto R (1998) Sex ratios, size distributions, and sexual dimorphism in the dioecious tree Ilex aquifolium (Aquifoliaceae). American Journal of Botany 85, 1602–1608.
Sex ratios, size distributions, and sexual dimorphism in the dioecious tree Ilex aquifolium (Aquifoliaceae).Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3MnhtlGjsQ%3D%3D&md5=7af046b54a5593b53accb3d68c82233eCAS | 21680320PubMed |

Oyama K, Dirzo R (1988) Biomass allocation in the dioecious tropical palm Chamaedorea tepejilote and its life history consequences. Plant Species Biology 3, 27–33.
Biomass allocation in the dioecious tropical palm Chamaedorea tepejilote and its life history consequences.Crossref | GoogleScholarGoogle Scholar |

Pickering CM, Arthur JM (2003) Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae). Austral Ecology 28, 566–574.
Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae).Crossref | GoogleScholarGoogle Scholar |

Popp JW, Reinartz JA (1988) Sexual dimorphism in biomass allocation and clonal growth of Xanthoxylum americanum. American Journal of Botany 75, 1732–1741.
Sexual dimorphism in biomass allocation and clonal growth of Xanthoxylum americanum.Crossref | GoogleScholarGoogle Scholar |

Porté A, Trichet P, Bert D, Loustau D (2002) Allometric relationships for branch and tree woody biomass of Maritime pine (Pinus pinaster Aït.). Forest Ecology and Management 158, 71–83.
Allometric relationships for branch and tree woody biomass of Maritime pine (Pinus pinaster Aït.).Crossref | GoogleScholarGoogle Scholar |

Scott SL, Aarssen LW (2013) Leaf size versus leaf number trade-off in dioecious angiosperms. Journal of Plant Ecology 6, 29–35.
Leaf size versus leaf number trade-off in dioecious angiosperms.Crossref | GoogleScholarGoogle Scholar |

Sudhakar P, Chattopadhyay G, Gangwar S, Ghosh J (2000) Effect of foliar application of Azotobacter, Azospirillum and Beijerinckia on leaf yield and quality of mulberry (Morus alba). The Journal of Agricultural Science 134, 227–234.
Effect of foliar application of Azotobacter, Azospirillum and Beijerinckia on leaf yield and quality of mulberry (Morus alba).Crossref | GoogleScholarGoogle Scholar |

Suzuki A (2000) Patterns of vegetative growth and reproduction in relation to branch orders: the plant as a spatially structured population. Trees 14, 329–333.
Patterns of vegetative growth and reproduction in relation to branch orders: the plant as a spatially structured population.Crossref | GoogleScholarGoogle Scholar |

Suzuki AA (2005) Reproductive investment in male and female Eurya japonica (Theaceae) at tree and branch levels. American Journal of Botany 92, 2003–2009.
Reproductive investment in male and female Eurya japonica (Theaceae) at tree and branch levels.Crossref | GoogleScholarGoogle Scholar | 21646119PubMed |

Swenson NG, Enquist BJ (2008) The relationship between stem and branch wood specific gravity and the ability of each measure to predict leaf area. American Journal of Botany 95, 516–519.
The relationship between stem and branch wood specific gravity and the ability of each measure to predict leaf area.Crossref | GoogleScholarGoogle Scholar | 21632377PubMed |

Tokár F (1985) Dry matter production of the aboveground biomass in relation to leaf area of the crowns in different types of young European chestnut stands. Folia Dendrologica 12, 161–175.

Xu X, Yang F, Xiao X, Zhang S, Korpelainen H, Li C (2008) Sex-specific responses of Populus cathayana to drought and elevated temperature. Plant, Cell & Environment 31, 850–860.
Sex-specific responses of Populus cathayana to drought and elevated temperature.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXnvVOmurs%3D&md5=08dea37f262e0aa7c5f883fbfa83597aCAS |

Zhang W, Jia X, Morris EC, Bai Y, Wang G (2012) Stem, branch and leaf biomass–density relationships in forest communities. Ecological Research 27, 819–825.
Stem, branch and leaf biomass–density relationships in forest communities.Crossref | GoogleScholarGoogle Scholar |

Zhang X, Zhang C, Zhao X (2014) Effect of sex ratio, habitat factors and neighborhood competition on stem growth in the dioecious tree Fraxinus mandshurica. Ecological Research 29, 309–317.
Effect of sex ratio, habitat factors and neighborhood competition on stem growth in the dioecious tree Fraxinus mandshurica.Crossref | GoogleScholarGoogle Scholar |

Zimmerman JK, Lechowicz MJ (1982) Response to moisture stress in male and female plants of Rumex acetosella L. (Polygonaceae). Oecologia 53, 305–309.
Response to moisture stress in male and female plants of Rumex acetosella L. (Polygonaceae).Crossref | GoogleScholarGoogle Scholar |