Updated Jan. 2009
Our lab studies the behavioral ecology of insect societies,
with a primary focus on the proximate and ultimate mechanisms
of honey bee queen behavior. In doing so, we attempt to address
questions of basic science that have practical relevance. Our
philosophy is to integrate a general understanding of bee biology
to help improve overall colony health and productivity; in an
era when the honey bee population is being severely impacted
by any number of factors, we feel that it is incumbent upon the
honey bee scientific community to become more proactive in asking
questions that address not just basic (long-term) or applied
(short-term) questions, but both.
1. Effects of genetic diversity on honey
bee colony phenotype
Our lab has a protracted history of investigations
into the adaptive benefits of intracolony genetic diversity.
Specifically, we have investigated the fitness advantages of
queen polyandry (mating with multiple males) and its consequences
on colony phenotype. Increased genetic diversity makes it less
likely that any particular trait is overly prevalent within a
colony. The fathers of the workers, a queen’s mates, carry
different alleles that vary with respect to many traits. Because
of the haplodiploid genome of honey bees, a queen that mates
once produces genetically similar workers that all carry the
same alleles from their father. If, by chance, his alleles are
unfavorable—for example, susceptible to a particular disease—then
all of the workers will be of low genetic quality and the colony
would be impacted severely under adverse conditions. A queen
that mates multiply, however, produces genetically diverse workers
that carry different alleles from their collective fathers. By
doing so, a queen reduces the risk that all of her worker offspring
will be of low genetic quality, increasing the probability that
the colony, as a whole, will be diverse enough to overcome a
wide range of ecological conditions. We have empirically tested
several non-mutually exclusive hypotheses that have shown the
adaptive benefits of genetic diversity to minimize the impact
of parasites and pathogens (Tarpy,
2003; Tarpy and
Seeley, 2006;
Seeley and Tarpy, 2007) and homozygosity at the complementary
sex determination (csd) locus (Tarpy
and Page, 2001, 2002).
2. Assessing the mating 'health' of honey bee queens
As an extension of this theoretical work, one of our newest
research paradigms is to investigate mating number of naturally
mated honey bee queens. There is both anecdotal and empirical
evidence to suggest that many queens produced in the U.S. may
not be adequately mated. Consequently, we are investigating
the current status of the “mating health” of honey
bee queens by looking at the physical quality, insemination
success, and mating numbers of a large sample of queen bees.
To measure mating number, we use PCR using microsatellite markers
to genotype workers in colonies and infer the mating numbers
of individual queens (Tarpy
and Page, 2000; Tarpy
and Nielsen, 2002; Tarpy
et al., 2004b). This is a large-scale project that
will enable us perform some very powerful investigations into
the evolution of extreme polyandry in honey bees, but it will
also ascertain whether or not a lack of genetic diversity within
honey bee colonies may partially explain the global ill-health
of commercial beehives.
3. Molecular mechanisms of honey bee mating
Together with Dr.
Christina Grozinger's lab (now at Penn State),
we have another research project to investigate the regulation
of honey bee reproduction by looking at the genomic and physiological
changes in queen bees during mating. We are using techniques
such as microarray gene-expression, classical behavioral observation,
and GC-MS to determine how virgin queens (who are receptive
to mating) transition to laying queens (who never mate again
in their entire lifetimes). Comparisons of virgin, partially
mated, and fully mated queens have revealed that physiological
changes in ovary maturation and pheromone profiles correlate
with gene-expression changes in the ovaries, while changes
in behavior correlate with gene-expression changes in the brain
(Kocher
et al., 2008).
We have also shown that queens inseminated with semen from
a single drone produce significantly different pheromone blends
that are less attractive than those from queens inseminated
with semen from 10 drones (Richard
et al., 2007).
4. Potential roles of nutrition and genotype on Colony
Collapse Disorder (CCD)
The accelerated losses of honey bee colonies in the fall and
winter of 2006-07 resulted in the bringing together of a working
group to explore the causes of a condition we have termed
colony collapse disorder, or CCD. This disorder is characterized
by the rapid loss of adult worker bees from affected colonies
with a resultant weak or dying colony with excess brood. The
CCD working group is focused on three main areas. First, the
potential influence of nutritional health on colonies,
either as a potential cause of CCD or an effect of another
factor. Linked to colony health is the genotype of colonies,
such that there may be potential genotype-environment interactions
that may be important in the incidence of the disorder. Second,
the possible influence of pathology on CCD-affected
colonies, either a more virulent version of an existing parasite
or potentially a currently unknown disease. Third, the possible
influence of environmental contaminants, either from
external sources (e.g., pesticides), internal sources (e.g.,
acaricides used to control varroa mites), or some combination
of sources. We are concentrating on the first of the above
potential causes, namely the role of nutritional stress and
genotype on the incidence and prevalence of CCD. We have quantified
whole-bee protein levels as a proxy for colony nutritional
stress. This measure will help us determine if CCD is associated
with poor colony nutrition, either as a potential cause (e.g.,
stressed colonies are more susceptible to pathogen attack)
or as a potential effect (e.g., collapsing colonies are less
able to acquire sufficient forage to maintain proper colony
health and function). Finding such an association will prompt
hypothesis testing of colony nutrition to determine cause versus
effect and possible mitigating management practices. We are
also currently genotyping colonies using microsatellite markers
to determine if intracolony genetic diversity has any bearing
on the disorder.
5. Competition among honey bee queens during the requeening
process
We also have an extended interest in the dramatic worker-queen
interactions during queen production in honey bee colonies.
When honey bee colonies rear new queens, they raise many but
then leave the queens fight to the death for only one to reclaim
the nest. The fitness discrepancy between “winning” and “losing” queens
cannot be greater: losing queens die (i.e., zero fitness),
whereas winning queens not only survive, they inherit an established,
successful nest with all the workers and inherent resources.
Clearly, natural selection ought to be acting very strongly
on any behavioral mechanism that governs which queen(s) survives
in order to bias the process in favor of certain queens. The
question is, how? We have investigated the potential roles
that worker may play in biasing the outcomes of competitive
duels (Tarpy and
Fletcher, 1998; Tarpy
et al., 2000; Gilley
et al., 2003; Gilley
and Tarpy, 2005), the physical advantages of individual
rival queens (Tarpy and Mayer, in review) including the fighting
tactics that they may use against each other (Tarpy
and Fletcher, 2003), and how selection may act on colonies
at the individual- and colony levels (sometimes in opposing
directions; Hatch
et al., 1999; Tarpy
and Gilley, 2004; Tarpy
et al., 2004a).
As such, this rich behavioral paradigm affords opportunities
to answer questions about levels of selection, kinship theory,
reproductive skew, and classic behavioral competition.
6. Non-chemical control of varroa mites
Varroa mites (Varroa
destructor) preferentially parasitize drone brood (developing
males) over worker brood because their development time is
longer compared to workers, enabling the female mite, or foundress,
an opportunity to produce a greater number of daughter offspring.
Because of this increase in reproductive potential, mites have
developed the ability to recognize and seek out drone brood
based on olfactory, gustatory, and mechanical cues. This differential
attraction to drone brood makes it a promising target for mite
control efforts, and previous studies have shown significant
control by removing drone frames and freezing them (along with
their mites). We are testing a new variation on this technique
by "rescuing" the drones after removing the mites from them.
This manipulation of drone brood could potentially benefit
the honey bees at the colony level as well as the population
level. The colony itself will benefit by decreasing infestation
of Varroa
destructor and
encouraging overall colony health and productivity. On a population
level, it is possible that surviving drones that live to sexual
maturity may possess some element of resistance to parasitism
by the Varroa mite. This research is being conducted by Holly
Wantuch, a Masters student in our program.
7. Mapping putative feral honey bee populations
Feral honey bee populations once
flourished in North America. Upon the arrival and establishment
of a parasitic brood mite, Varroa destructor, feral
honey bee populations were largely decimated. While it has
been hypothesized that these once-thriving feral populations
served as a genetic reservoir of diversity for managed honey
bee populations in the United States, no empirical evidence
is available to support this claim, to date. In response to
growing concerns about levels of genetic diversity in managed
honey bee populations in the United States, plans to import
additional honey bee germplasm are currently being addressed.
This leads to the question of the status of feral populations.
Research and anecdotal evidence suggest that the feral population
may be recovering (see SaveTheHives.com for a current map).
We are currently investigating the status of feral honey bee
populations in the U.S. by re-sampling non-managed honey bee
populations in the southeastern United States to determine
its genetic composition and to finally address whether feral
populations are truly "survivor stock" or simply recent "escaped
swarms". Results from this study will reveal the amount of
genetic diversity available within U.S. honey bee populations
and will allow us to determine the extent of genetic loss.
This research is being conducted by Dr.
Debbie Delaney, a USDA
Postdoctoral Fellow in our program.
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