Stem tissue mass density is linked to growth
and resistance to a stem-boring insect in
Alternanthera philoxeroides
XIAOYUN PAN,* XIN JIA,* JING ZENG,* ALEJANDRO SOSA,† BO LI* and JIAKUAN CHEN*
*Coastal Ecosystems Research Station of the Yangtze River Estuary, Ministry of Education Key Laboratory for Biodiversity
Science and Ecological Engineering, Institute of Biodiversity Science, Fudan University, Shanghai, China; and †South American
Biological Control Laboratory, USDA-ARS, Hurlingham-Buenos Aires, Argentina
Abstract
To investigate how stem anatomical structure is linked to growth and resistance to
stem-boring insects in a herbaceous species, six populations of alligatorweed (Alternan-
thera philoxeroides) were grown in a common garden. Stem growth rate (GR) of A.
philoxeroides and pupation rate as an estimate of resistance to a stem-boring insect
(Agasicles hygrophila) were quantified. Stem tissue mass density (TMD) was measured
and stem anatomical traits were analysed on cross-sectional areas (CSA). Stem TMD was
positively correlated with resistance (i.e. negatively correlated with pupation rate) and
negatively correlated with GR. Stem cortex CSA (%) and vascular bundle (VB) density
(no./mm
2
) were positively related to stem TMD and negatively related to pupation rate.
The GR was positively related to VB CSA (%) and negatively related to VB density. These
results suggest that stem TMD, which results from a high fraction in cortex CSA and high
VB density, is a key determinant of resistance to a stem-boring specialist in A. philox-
eroides. The high resistance of plants with higher stem TMD may partially impose a cost
to plant growth.
Keywords: growth rate, herbivore resistance, stem anatomy, stem tissue mass density, stem-
boring insect.
Received 1 January 2010; accepted 29 August 2010
Introduction
Recently, plant tissue mass density (TMD; the ratio of dry
mass to fresh volume) has attracted considerable attention
in comparative ecology (Westoby & Wright 2006; Chave
et al. 2009; Poorter et al. 2010). It can be seen as a syndrome
of traits involving many potentially competing functions,
such as carbon gain and growth potential, mechanical
strength, efficiency and safety of hydraulic transport, and
resistance to herbivory (Sperry et al. 2008; Chave et al.
2009; Poorter et al. 2010). Wood density partially under-
lies, for example, the growth–survival trade-off that is
observed among woody plants (Poorter et al. 2010). Fast-
growing species with low-density tissues are character-
ized by leaves in which a high fraction of volume is
occupied by mesophyll, the assimilatory tissue (van
Arendonk & Poorter 1994). In contrast, slow-growing
species produce high-density tissues, which favour both
organ persistence and conservation of resources (Gleeson
& Tilman 1994), and resistance to herbivorous insects
(Coley et al. 1985). Although the relationship between
TMD, anatomy and whole-plant functioning, such as
growth, height and safety of hydraulic transport and
mechanical stability, is widely recognized for stems, there
is little information about such a relationship among stem
TMD, growth and resistance to stem-boring insects.
The aim of the present study was to investigate whether
two key traits in whole-plant functioning, that is, growth
rate and resistance to a stem-boring specialist, are related to
stem TMD and stem anatomy in a herbaceous species,
Alternanthera philoxeroides (Mart.) Griseb. (alligatorweed;
Amaranthaceae). Previous studies have shown that under
controlled conditions this species differs substantially in
growth rate and resistance (Jia 2008). Six populations of A.
philoxeroides were grown in a common garden. Anatomical
Correspondence: Xiaoyun Pan
Email: xypan@fudan.edu.cn
Plant Species Biology (2011) 26, 58–65 doi: 10.1111/j.1442-1984.2010.00307.x
© 2010 The Authors
Journal compilation © 2010 The Society for the Study of Species Biology