Plant Ecology & Diversity, 2013
http://dx.doi.org/10.1080/17550874.2013.777133
Divergent seed production responses of white and blue flowers of Gentiana leucomelaena
(Gentianaceae) to warming and watering
Junpeng Mu
a,b
, Youhong Peng
c
and Kechang Niu
a
*
a
Department of Biology, Nanjing University, Nanjing, China;
b
Ecological Security and Protection Key Laboratory of Sichuan Province,
Mianyang Normal University, Mianyang, China;
c
Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan
Province, Chinese Academy of Sciences, Chengdu, China
(Received 14 April 2012; final version received 13 February 2013)
Background: The adaptation, evolution and function of flower colour diversity in response to changing environments are one
of the oldest puzzles in plant ecology. It is logical that comparative studies on fitness of flower colour would be the most
meaningful if they were conducted on the same genotype.
Aims: We used Gentiana leucomelaena, which produces both white and blue flowers on different tillers of the same individual,
as a model species to test adaption and fitness differences of contrasting flowers in contrasting environments.
Methods: We examined seed production by white and blue flowers in response to increased temperatures (28
◦
C/2
◦
C vs.
12
◦
C/2
◦
C; 12 h light/12 h dark) in a climate chamber study and to watering in the field (+1.5 l m
-2
d
-1
vs. control with no
supplementary irrigation).
Results: For white and blue flowers warming decreased but watering increased seed number. Increased temperature and
watering increased the size of seeds from white flowers, but neither warming nor watering significantly changed seed size of
blue flowers. Seed size was significantly negatively correlated with seed number in the temperature treatments but positively
correlated in watering treatments. The positive correlation was strong in white flowers, but the negative correlation was
stronger in blue than in white flowers.
Conclusions: Water availability and low temperature confer an advantage to white flowers, while warming and dry habitats
favour blue flowers. These divergent responses may influence total plant fitness and thus help explain the adaptive value and
evolution of flower colour diversity.
Keywords: adaptation; flower diversity; reproductive strategy; seed number; seed size
Introduction
The evolution and function of flower diversity (e.g. colour,
size, shape) is one of the oldest puzzles in plant ecol-
ogy, and there has been much research on the different
exogenous and endogenous determinants of flower diversity
(Darwin 1859; Stebbins 1974; Galen 1999; Dormont et al.
2010). Previous studies on flower colour diversity often
have focused on the effect of biotic factors on the evolu-
tion of the colour diversity, e.g. attracting insect pollinators
(Ida and Kudo 2003; Chittka and Raine 2006; Kudo et al.
2007). Recently, with global climate change, an increasing
number of studies have been concerned with the rapid adap-
tation and evolution of flower colour diversity to changing
abiotic factors (Orenshamir and Levinissim 1997; Grace
and Logan 2001; Steyn et al. 2002). However, there is
a scarcity of experimental studies on the relative fitness
of different flower colours under varying environmental
conditions. The results of such studies would help us to bet-
ter understand the evolution and function of flower colour
diversity.
Previous studies on the function of flower colour diver-
sity often have compared the fitness of individuals in
different species or populations that had contrasting flower
colours; however, due to genetic differences it is difficult to
*Corresponding author. Email: kechangniu@nju.edu.cn
determine the functional role of flower colour as a determi-
nant of fitness. Theoretically, comparing the fitness between
phenotypes within the same genotype (e.g. between tillers
of the same individual) with contrasting coloured flowers
is better than comparing genetically different members of
a species with different colours. In plants with the same
genotype, genetic variation would not be a constraining fac-
tor in determining the fitness of different flower colours.
However, there are very few studies comparing the adap-
tation and fitness of contrasting flowers among individuals
within a population or among tillers in the same individ-
ual. Flower polymorphisms are common in nature, e.g. both
blue and white flowers are produced by Linanthus parryae
(A. Gray) Greene, a California desert annual (Schemske
and Bierzychudek 2007). Examples of species with tillers
on the same plant that produce different colours of flow-
ers are not common, but one example of such a species
is Gentiana leucomelaena Maximowicz ex Kusnezow. In
this species some tillers produce white flowers and others
produce blue flowers (Figure 1). The use of G. leucome-
laena as our experimental organism allowed us to control
for the effect of genetic variation in comparative studies on
the fitness (e.g. seed number and seed size) of flower colour
and on responses of contrasting flower colours to climate
© 2013 Botanical Society of Scotland and Taylor & Francis
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