Functional
Ecology 2000
14, 215 – 225
© 2000 British
Ecological Society
215
Blackwell Science, Ltd
Effects of herbivory on internal ethylene production and
sex expression in Cucurbita texana
G. A. KRUPNICK,* G. AVILA, K. M. BROWN and A. G. STEPHENSON
Department of Biology, 208 Mueller Laboratory, The Pennsylvania State University, University Park, PA 16802, USA
Summary
1. Field-grown Cucurbita texana was used in a study to determine if stress ethylene
from damaged leaves promotes changes in sex expression.
2. Internal ethylene concentrations were experimentally enhanced by inserting an
ethylene-filled syringe into the hollow chamber of an internode seven nodes from the
growing tip of a branch. Branches enhanced with ethylene produced significantly
more pistillate buds than control branches and experienced greater bud abortion
later in floral development.
3. The timing and magnitude of Diabrotica beetle leaf damage was simulated using a
paper-hole puncher (15% removed from all leaves). Simulated herbivory was applied
to new growth every 3 days throughout the growing season.
4. Endogenous ethylene concentrations recovered from the internode adjacent to the
growing tip were significantly greater in damaged plants than undamaged plants 1 day
after damage to new growth during the second and third week of the experiment, and
2 days after damage during the fourth and fifth week. By the sixth week, no significant
differences were evident.
5. Damaged plants produced fewer pistillate buds than undamaged plants, indicat-
ing that stress ethylene from simulated herbivory does not induce femaleness. A
second messenger that suppresses pistillate bud production may be produced during
damage.
Key-words: Insect damage, monoecious, phytohormones, reproductive biology, sink–source interactions
Functional Ecology (2000) 14, 215 –225
Introduction
Like other phytohormones, the gaseous hormone
ethylene has multiple functions in plant growth and
development (Abeles, Morgan & Saltveit 1992). For
instance, ethylene is involved in the regulation of fruit
ripening and sex expression, and as a response to her-
bivore damage (Abeles et al. 1992). In a previous study,
we found a link between two of these functions in
Cucurbita texana (Scheele) A. Gray (Cucurbitaceae) –
internal endogenous ethylene concentrations climb dur-
ing fruit maturation and ethylene produced from mature
fruits correlates positively with pistillate flower produc-
tion at the tip of growing branches (Krupnick, Brown
& Stephenson 1999). In short, the final stage of fruit
development corresponds with the differentiation of
additional pistillate floral buds by way of a hormonal
signal. While the timing between fruit ripening and
pistillate flower bud production may be an appro-
priate way to separate competing resource sinks (i.e.
fruit) adequately, stress-induced ethylene may be an
inappropriate regulator for pistillate flower production
because leaf herbivory may reduce the acquisition of
resources and the amount of stored resources needed
for successful fruit development and, in some circum-
stances, it may divert resources away from reproduc-
tion and towards inducible defenses (Baldwin, Sims
& Kean 1990; Karban & Baldwin 1997). Nevertheless,
if herbivory increases ethylene concentrations, stress-
produced ethylene may also regulate the differentiation
of additional floral buds. In this study, we examine
whether the sex expression of C. texana responds to
ethylene induced by herbivory in the same way that it
responds to endogenously produced ethylene.
Several studies have shown that ethylene regulates
sex expression in many monoecious species including
members of the Cucurbitaceae (Rudich 1990; for review).
Studies have used exogenous growth regulators (e.g.
ethephon, as an ethylene-releasing agent), ethylene
biosynthesis inhibitors (e.g. aminoethoxyvinyl glycine)
and ethylene action inhibitors (e.g. silver nitrate and
silver thiosulphate) to show that ethylene promotes
*Present address: Department of Botany, NMNH, Smithso-
nian Institution, Washington, DC 20560 –0166, USA.