Magazine

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variability are well-studied. On the other half remaining on a

Quick guide the island of Trinidad, lowland non-recombining Y. And this Y
areas of streams have a rich chromosome, similar in size to
community of predatory fish, the X, is itself variable between
Guppies while further upstream, barrier male lineages. In parallel with the
waterfalls have allowed guppy amplification of male coloration
Felix Breden populations to evolve without genes, genes encoding
large predatory fish. Through photoreceptive opsin proteins
various approaches, it has been have been duplicated and
What are guppies? Guppies shown that behaviors, life history exhibit a high degree of adaptive
(Poecilia reticulata) are small and morphology all respond diversification.
tropical fish, native to the coastal evolutionarily to the strength of
streams of northeast South predation. For instance, males What is the guppy’s potential
America. They owe their name tend to be brighter and more for biomedical research?
to Robert John Lechmere Guppy conspicuous in areas of low With their fast generation times
who introduced them into the predation. Thus, the guppy is one and ease of maintenance,
aquarium trade. Guppies have of the few systems in which the guppies are a valuable resource
since become very popular dominant selective force can be for biomedical research. For
aquarium fish, known for colorful so clearly identified. Manipulative instance, David Reznick and
males and live-bearing females. experiments within and between colleagues are exploiting
But guppies are also one of streams have shown that guppy population differences in
the premier model systems for populations can evolve extremely life history as a model for
the study of ecology, evolution, rapidly, at speeds similar to those understanding the forces
genetics, and sexual selection. observed in artificial selection shaping variation in aging, and
The guppy is now poised to experiments in the lab. in our lab we are studying the
become an important model for guppy mutant curveback as a
functional genomics of adaptive What about guppy genetics? so far unique model for familial
characters. The genetics underlying these idiopathic scoliosis.
characters are also unique.
What is so special about Beginning in the 1920s, Winge Guppy genomics? Detlef Weigel,
them? Guppies are extremely and colleagues showed that Christine Dreyer and colleagues
variable both phenotypically over 40 loci contribute to male in Tübingen, Germany, are
and genetically. Sexually mature coloration. Intriguingly, all of currently constructing a genetic
males exhibit an amazing array these loci are linked to the linkage map to associate variable
of differently colored spots sex chromosomes, with about phenotypic traits with genes of
and stripes, such that every half of the alleles recombining known function. The mapping
male almost seems unique between the X chromosome crosses will include highly
(Figure 1), making the guppy and the Y chromosome in a differentiated and long- studied
one of the most polymorphic pseudoautosomal fashion, and natural populations. Our lab has
vertebrates known. Although
the females do not show such
coloration, they vary in terms
of their preferred mates both
within and between populations,
making the guppy a powerful
system for studying sexual
selection. Like other members
of the family Poeciliidae, such
as swordtails and mollies,
female guppies are live-bearing.
Males fertilize the eggs using
a stick-like modified anal fin,
the so-called gonopodium.
Guppies are ovoviviparous, i.e.
the eggs develop inside the
mother. However, also viviparity
with placental nutrition has
evolved several times within the
Poecilliidae.

What have guppies taught Figure 1. Guppy diversity.

us about evolution? The An array of males retrieved from a single population of the Cumaná guppy reveals the
guppy evolves quickly and the diversity of male coloration in these fish, which is controlled by more than 40 sex-linked
selective factors that shape this loci. Photo used with permission from Alexander and Breden (2004).
Current Biology Vol 16 No 20
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been using microsatellite markers For plants, associations with

developed in the closely  related
Primer fungi and bacteria were key
sword-tail Xiphophorus innovations in the colonization
maculatus, a known model for of land and of specific habitats.
the genetics of melanoma, to Symbiosis Eukaryote-associated microbes act
quickly build a set of linked as metabolic partners for accessing
markers in the guppy. Thus, the Nancy A. Moran limiting nutrients and also as
large group of biologists studying protectors, producing toxins that
the family Poeciliidae is joining Microorganisms arose and ward off herbivores or pathogens.
efforts in establishing genomic diversified before the appearance Similar associations have arisen
resources for this valuable group of large multicellular organisms. with animals, allowing colonization
of species. The bodies of the latter provided of diverse niches, such as the
new potential habitats for specialized feeding on plant or
What is in store for the microbes — habitats that were animal tissues, and the use of deep
guppy? Variation is essential persistent, stable (from a microbial ocean hydrothermal vent habitats.
to understanding the genetics perspective) and nutrient-rich. Often, the associations are
underlying biological processes. As a result, large organisms, persistent for the hosts, frequently
The guppy provides a huge from oaks to humans, have being transmitted vertically across
amount of natural variation been continuously enmeshed generations, from mother to
that can be studied genetically. in complex interactions with progeny. The organisms involved
This variation has not been microorganisms during their in a symbiosis may be sufficiently
produced by mutagenesis evolution. Biologists have paid fused that they cannot live apart or
screens, but instead has been most attention to associations be recognized as distinct entities
shaped by natural selection. with microbes that are pathogenic; without close scrutiny.
Thus, combined with emerging typically, microbial infection has
genomic resources, the been viewed as deleterious, or Recent growth of research on
guppy is an ideal organism for at best irrelevant, to vigor and symbiotic microorganisms
understanding the evolutionary reproduction. But the last 15 Before biological research
genetics and molecular basis of years have witnessed increased became focused on genetic
adaptation. appreciation of interactions that model organisms, greater
benefit the host as well as the attention was paid to the study
Where can I find out more? microbe. These interactions are of symbiosis. Microscopy was
Alexander, H. and Breden, F. (2004). Sexual
isolation and extreme morphological
loosely grouped under the term used extensively to discover and
divergence in the Cumaná guppy: a ‘symbiosis’ and the microbial elucidate symbioses before the
possible case of incipient speciation. J. partners called ‘symbionts’. middle of the 20th century. The
Evol. Biol. 17, 1238–1254.
Endler, J.A. (1994). Multiple-trait coevolution Although the term ‘symbiont’ is reasons for a period of relative
and environmental gradients in guppies. typically applied to mutualistic neglect, until about 1990, are
Trends Ecol. Evol. 10, 22–29.
Houde, A.E. (1997). Sex, Color, and Mate
microorganisms, it is often used to complex. One contributing factor
Choice in Guppies. (Princeton, New include associates for which the has been the unsuitability of most
Jersey: Princeton University Press). full spectrum of effects on hosts is symbiotic partners as models for
Lindholm, A., and Breden, F. (2002). Sexual
selection and sex chromosomes not known. laboratory studies of genetics
in poeciliid fishes. Am. Nat. 160, Symbiosis is ubiquitous in and development. The organisms
S214–S224.
Magurran, A.E. (2005). Evolutionary Ecology:
terrestrial, freshwater and marine that are easiest to grow and
The Trinidadian Guppy. (Oxford, UK: communities. It has played a key study in the laboratory, such as
Oxford University Press). role in the emergence of major Escherichia coli, Arabidopsis
Olendorf, R., Rodd, F.H., Punzalan, D.,
Houde, A.E., Hurt, C., Reznick, D.N., life forms on Earth and in the thaliana, Drosophila melanogaster
and Hughes, K.A. (2006). Frequency- generation of biological diversity. and Mus domesticus, are weedy
dependent survival in natural guppy
populations. Nature 441, 633–636.
Numerous authors have pointed species adapted to show fast
Reznick, D.N., Shaw, F.H., Rodd, F.H., and out that Darwin did not emphasize growth in temporary niches.
Shaw, R.G. (1997). Evaluation of the rate symbiosis as an evolutionary These species have lifestyles
of evolution in natural populations of
guppies (Poecilia reticulata). Science mechanism; its role is, of course, relatively free of complex
275, 1934–1937. not inconsistent with Darwin’s interdependencies with other
Reznick, D.N., Mateos, M., and Springer,
M.S. (2002). Independent origins and
main ideas, but appreciation species. But most microorganisms
rapid evolution of the placenta in the of symbiosis as a source of in natural communities are likely
fish genus Poeciliopsis. Science 298, evolutionary novelty has developed to have obligate dependencies
1018–1020.
Reznick, D., Bryant, M., and Holmes, D. relatively recently. Its importance on other species (often other
(2005). The evolution of senescence and is perhaps most strikingly microbes), explaining why 99%
post-reproductive lifespan in guppies
(Poecilia reticulata) PLoS Biol. 4, e7.
illustrated by the symbiotic of microorganisms are difficult or
events that led to the evolution of impossible to culture. Similarly,
eukaryotic organelles — plastids most symbionts of plants and
and mitochondria — from animals cannot be readily cultured
Department of Biological Sciences,
Simon Fraser University, Burnaby, cyanobacterial and independently of hosts, precluding
British Columbia, Canada. alpha- Proteobacterial most conventional microbiological
E-mail: breden@sfu.ca ancestors, respectively. analyses.