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The varroa mite ( Varroa destructor Anderson and Trueman)

Reproductive biology of Varroa

is an ecto-parasite of western honey bee ( Apis mellifera ) and is

distributed worldwide. apis mellifera colonies usually die
within two to three years after of mite infestation, if untreated,
colonies of wild bees in

the U.S. .

were almost completely


exterminated by this mite around 1995 (circa 1987).

Anecdotal evidence that bees could be increasingly aggressive
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with mites in recent years (genetics of resistant mite strains, but

no a systematic study for this claim. The varroa mite is
currently the most serious pest of honey bees worldwide.


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Understanding the reproductive biology of the Varroa mite will

allow us to better manage this important pest in our apiaries.

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The life cycle of Varroa

The life cycle of the mite Varroa has two stages. During the
phoretic stage are mites on adult bees or drones, while feeding
on the blood (hemolymph) of bees, usually inter-membrane
segmental in the abdomen. phoretic stage lasts about 5-11 days
until the cell enters a larva bee or drone. course, the mites are






forced to stay in phoretic state if no breeding, and this can last

5-6 months in cold climates. Mites change hosts (jump from
one bee to another) often and this contributes to the spread of
various viruses, infecting all parasitic bees during feeding.
Mites experience increased mortality during the phoretic stage,
and fall to the bottom of the hive. However, the total of these


fallen mites are less than 20% of the population. therefore

using a bottom plate reduces but does not eliminate the use of
chemicals to manage varroa. 's phoretic stage is important for
mites to spread to other colonies for foraging or drift another
colony, or, finally, by the plundering of weaker colonies. In the
latter situation, we are actually selecting mites with high

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virulence, because while isolated hives mites in a colony will die

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with its host (due to the low probability of being found by a

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neighboring colony), while in an apiary this behavior plunder

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weaker hives by disease or by the same varroa is guaranteed,

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ensuring successful transfer of mites to other sick and weak

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hives of apiary colony, in which the cycle is repeated again.

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The other step is the reproductive stage, and only during this
time may increase mite population. This occurs only in the
brood cells of workers and drones. 's to expensive mature female
bee engage when they emerge from the cell. The varroa mite
invades a host cell just before the cell is capped. Once inside,
hidden in the larval food in a prone position (view from the top
of the cell). mites have special appendages called "peretrimes"
(essentially as snorkels) that help you breathe when it introduced
into the larval food. Shortly after the larva inside a cocoon
built and then will become a prepupa. mite not be fed to about
five hours after the larva completes its cocoon and put the first
egg 7 0 hours after operculado cell. The first egg is not fertilized
and becomes a male. , this mechanism the same as in the bees,
since both species have what is called the mechanism
"haplodiploidy" sex determination, so that the males are haploid
(which has no parent) and females are diploid (with both
parents). Thereafter, approximately every thirty hours, the
female mite lays an egg. mite If the mother is not properly
fertilized, then the offspring will be male only when emerge. to a





total of five (as pupae workers) or six eggs (about drone pupae.)
However, as will emerge honeybees eleven days after capping,
and drones fourteen days a mite daughter has six days to
mature, most of these eggs do not have time to develop into
adults (6 + 70 hour delay in egg laying + 1 day for the first
egg as male = 10 days, leaving only a daughter mature).
Males and females that have not been fully developed, all die
shortly after due to dehydration after opening a cell (after
emergence of the bee, or be opened by bees with hygienic behavior
.) Therefore, only the mites mature and darker color female,
but not most of the stadiums and males mature nymphs are seen
by most beekeepers. males mate with a female several times to
result in a total of about 35 sperm stored within the female

Varroa has "faecal sites" in the cell, where they deposit their
feces, which are white in color due to a high concentration (~
95%) of guanine. For some unknown reason, any mite defecates
in the pupa is directly sterile.
Methods for studying mite reproduction
There are two methods for the study of mite reproduction. A
method is simply uncover worker cells or drone in the colonies
and the percentage of mites that reproduce (fertility), or the
number of offspring (fecundity) is determined mites. This
method provides information about what happens in the wild,
but the information obtained is limited because it is not
controlled or manipulated. Another method is to perform
manipulations, either mites, then artificially introduced mites in
cells and waiting 9-10 days to determine fertility and





fecundity. Combs can be reintroduced into a colony (at the risk
of elimination due to bee hygienic behavior), or incubated in a
laboratory. The basic steps for this process are as follows.


's Mites come from brood cells or a colony with a high

number of mites. mites now routinely harvested using the

method of powdered sugar, then the sugar is cleaned using a
damp brush.
2). Obtain brood cells that have a limit (6 hours). mites will
not play if introduced into cells that have been capped after 14
hours. Ideally, the cells must be sealed within the last . six
hours Scientists often use a piece of transparency to assign
offspring; marking being introduced (with holes in the lid),
waiting for six hours, and labeled again. Those cells which have
a hole at the top, were totally obscured then the cells are
needed. We ensure mark the two holes (fixing transparency on
Honeycomb) because bees can seal small holes with wax, and the
reference is lost.
3). slightly Open cell using a fine scalpel, insect pin, or a pair
of fine forceps, and carefully introduce a mite small slit using a
horsehair brush or fine.
4). Push the wax level it, and sealed with melted beeswax with
a paintbrush.
5). Maintain upright frame at all times, and the relative
humidity to 50%, and a temperature of 32-35 C.
. 6) Check the frame daily for signs of larvae of wax moth,
which may destroy the data; not place the flat side of the frame
on a surface, hold it upright during this process.
7). Finally, the ninth (day of transfer of mites, designated as
the first day), you can open the cells and count and record the
number of males, daughters mites (mature deutonymphs and
protonymphs) that are there.





During Stage Host phoretic

Whether varroa mites can choose multiple hosts in the phoretic
stage previously studied, using reared bees in cages and petri
dishes. The mites prefer nurse bees when they appeared in a
petri dish with pecoreadotas bees and nurses. mites also
transferred more frequently to young bees to old bees. This
discrimination Varroa later shown to be related to the repellent
effect of geraniol, a component of the pheromone gland
Nasonov, which is high in foraging bees. however, was not clear
whether mites show the same preference under the same
conditions in a hive in the field. A study showed that nurse
bees had a higher percentage of newly hatched mites, but no
differences between nurses and foragers bees were found.
Another study found that nurses were the most was preferred,
but the experiment was conducted in a colony (ie, not
replicated). was mites distribution among nurses age day study
(5-11 days old at the marked bees) and foragers (unknown age,
but the average age of foraging bees should be greater than 21
days in a typical colony), and found a clear preference for
farrowing. therefore mites do show the same preference for
nurses, even environment of the colony.





Why the state phoretic?

Scientists wonder why the mites go through a phoretic stage,
because they experience a high mortality rate during this period.
Under laboratory conditions, the varroa can successfully
reproduce without phoretic stage. Ie the mites transferred
immediately after the appearance of the bee to another cell of
breeding point opercurlarse with a maximum of seven cycles. the
average total number of children (including males) was four
during the first four to five cycles. This seemed greater that in
our experiment. Yet closer inspection, had mites had no

(21.7%), and

that this was most likely not included in

the calculation. So the actual fertility was

3.17 ,

4 * (1 to 0,217) =

which also included males. After subtracting the males

(which were about two per mother, instead of one, curiously, in

his study, De Ruijter, 1987), we have 1.17 female offspring
per mother. This would be slightly less than the 1.6 female
offspring of mother mites observed in fed newly hatched bees.
therefore mites phoretic not experience stage have lower fertility,
on especially compared to those hosted by younger nurses. Our
recent study showed that mites prefer nurse bees, perhaps not
only because of its proximity to the larvae (which feed the nurses
inspected and frequently), but also because the nurses provide
additional nutrition for playback on your phoretic phase. Our
study found that mites artificially fed in the nurses had the
greatest number of offspring, followed by the collectors, and fed
newly hatched bees, which have fewer offspring. Moreover,
when the fecundity of mites was compared hosted by bees of
different ages, a significant negative relationship between mite
fertility and the age of the nurses was found





Differences in mite reproduction is important for resistance

against mites
The varroa can be played on both worker bees and drone brood
in Apis mellifera , but played exclusively on drone brood in A.
cerana , the original host. many factors such as hygienic behavior
(removal of phoretic mites during stage adults), hygienic
behavior (removing mites pupae during the reproductive stage),
the duration of the state of farming and the attractiveness of
farming contribute to the varroa tolerance (reviewed by Bchler,
1994). However, I think the reduced reproduction (including
both reduced fertility and fecundity) in offspring of workers is
the most important for the strength of the bee against factor
Varroa . This is due to the infertility of the Varroa in breeding
of workers, correlates well with the degree of tolerance bee mite.
, for example, A. cerana is highly tolerant to the mite and
causes 100% of infertility mite in the brood, the Africanized bee
( scutellata A. mellifera ) is moderately tolerant of infertility
40%, while A. mellifera . is the least tolerant of lower fertility
rate (10-20%) in the brood. strain in European bees was
artificially selected to be tolerant of mites, mite infertility has
played the most important role in reducing the mite population,
while other factors (such as hygienic behavior, and duration of
operculado) are not as important (and Hoopingarner Harbo,
1997). Though we now know that the


Original (removing mite reproduction) feature is actually due to

"VSH" (hygienic


VSH can

be considered a special

lower causing playback feature, due to the interruption of the

reproductive cycle of mites especially because bees do not open
the cells containing mites do not reproduce, but that target those
fertile mites.





Factors affecting mite reproduction

Effect of breed breeding
It has been long known that varroa mites prefer drone brood
over worker brood, in a ratio of nine to one. Ie if an equal
number of cells available, the drone brood would house nine
times the amount of mites worker brood. Natural selection
undoubtedly favors drones mites preferred because drones
raising has a capping over period, which allows daughter more
mature mites. Indeed, Martin (1994, 1995 ) calculates the
effective rate of reproduction (ie, the number of viable / ripe
daughters by the invasion of the mother) and 1.3 to 1.45 in one
infested breeding working, while for drone brood was 2.2-2.6.
In A. mellifera , the transfer of mites for drone brood mite
reproduction always decreases, while the transfer of mites to the
workers drone brood increased the rate of reproduction. Queen
larvae are a dead end for the invasion of mites, because the
queens emerge at 16 days, five days faster than a worker,
leaving a daughter mites without time to mature. 's royal cells
prevents varroa apparently due to some chemical smell of royal





Effect of host species

Mites transfer across different species suggests that host species
also affects mite reproduction. When mites A. cerana
introduced in A.mellifera


in worker brood, only 10% of the

mites reproduced , while 80% of mites

A. mellifera mites were

transferred to A. cerana were reproduced in the brood. In our

study, Varroa destructor , is (> 90% play) played equally
well, whether transferred to A. mellifera or A. cerana , both
drone and worker. By contrast, V. destructor from A. cerana
only reproduced in A. cerana in drone (83% reproduced, N =
62), These results suggest mites in the two species of honey bees
are different: mites A. cerana

is refrain from reproducing in

worker brood of the same species, and mites of A. mellifera are

well reproduced in the capped brood regardless of the host
species. seems that only the haplotype V.destructor had a genetic
change that allowed him to play on any drone or worker brood
of A. mellifera , thus allowing it to build up to harmful levels
for bees. In China found no harmful levels of V. destructor in
A. cerana - in fact, in most locations, mites they could not be
found. When we found was the haplotype of Vietnam, which are
not published in the brood of A. cerana .'s not clear why the
Korean haplotype of V. destructor does not damage A. cerana ,
and that can be played on both the worker and drone brood in
transfer experiments. However, you may not reproduce in
worker brood under natural conditions. therefore transfer
experiments should be complemented with the observation in
natural conditions for a complete





Effect of cell size

Partly because the mites reproduce better in drone brood in
worker brood, people tend to think that with smaller cells mite
reproduction is reduced. However, two recent studies also show
that there was no difference in the population of mites between
colonies (Ellis et al., 2009) using the "small cells" (4.8 to 4.9
mm in diameter) of cells (5.2 to 5.4 mm), small cells actually
had a significantly higher mite population (Berry et al., 2010).
Unfortunately, none of these recent studies determined fertility
or fertility of the mites in the two cell types. Previous studies
were conflicting. Taylor et al. (2007) found that the cell size
did not affect the reproductive success of V. destructor , but had
more mites invaded cells taken from the base of 4.8 mm. Yet
Piccirillo and De Jong (2003) and Maggi et al. (2010)
found that the rate of invasion of mites increased positively and
linearly with the width of the cells of worker and drone brood,
probably due to breeding takes place in the large cells are more
views of the nurse, which increases the possibility of invasion.
Maggi et al. (2010) also found that the percentage of fertile
mites was lower in smaller cells. One previous study
(Goncalves, 1995 ) showed that AHB, larger cells have a
higher rate of invasion, and also had the highest effective
fertility mites. Our own study suggests that cells that are too
large also reduce mite reproduction (Zhou et al. 2001). A
study is to determine the mechanisms of the reason that the
varroa can not be played in worker brood of A. cerana , we
discovered by chance that both A. cerana and A. mellifera


queens laid eggs . of workers in the cells of drones in the fall

We took advantage of this, and reproductive performance of
mites was compared in two armies. workers reared in cells of
workers or workers reared in drone cells in 2001, both fertility
and fertility of the two groups were significantly different. 's not
clear why the mites reproduce less staying in larger cells. One
possibility is that the workers reared in drone cells are fed a
different diet by nurse (A study workers showed that raised in
drone cells were heavier

and had the largest

ovaries, suggesting a different diet or more nutrients). A

second possibility is that the workers larger spin cocoons at the





drone cells mites detect the extra space, and this affects their

Effect of humidity
Kraus and Velthuis (1997) wondered why varroa mites were
not such a big problem in the tropics (besides the fact that most
of the bees were from Africa), and tested in the laboratory to
see if moisture high relative inhibit mite reproduction.
individual mites were transferred into fresh artificial
operculadas cells, and after raising an incubator maintained.
When the relative humidity was set at 59-68%, average 53%
of the mites was (N = 174 mites) of offspring; under 79-85%
relative humidity, only 2% (N = 127) were reproduced mites.
The difference in mite fertility was highly significant. With
75% (instead of 50%), very few mites reproduce , with 80%
moisture varroa mite population will never increase to a harmful





Effect of Movement Honeycomb

Apart from where they defecate, varroa mites are also very
picky about where they feed. 's mother teaches her young a
feeding site on the pupa in particular (between the pair of rear
legs on the ventral side of the abdomen), and then leads back to
the site of defecation. therefore any rotation of the panels will
produce a movement of the host pupa? perhaps cause
disorientation mites.

exists a beehive with rotating frame

[broodnest "was invented (and patented) in Hungary. The hive

body is round frames and rotated ten degrees per hour,
completing a circle in 36 hours. This powered by a 12 volt

Loa Varroa mites are not able to reproduce because

constant turnover of cells. however, one study (Aumeier et al.,

2006) of rotating panels in mite reproduction for three years
found no evidence that it worked. 's Daily rotation or shaking
brood cells or "affect fertility (93-100%), and fertility (2.63.0) of reproductive mites or pup mortality of mites in brood
cells." This is a bit surprising because thought before applying
for the patent, the inventor should have obtained data showing
that rotation affected mite reproduction? However, the study
reported that the actual cells to form a swarm withdrew due to
rotation, so Estala hive only serves to prevent swarming.





Effect of Age, hormones, pheromones, and Genes

Varroa mites that have been artificially introduced into brood
cells that have been sealed 14 hours before the introduction of
the varroa will not play. Among the mites were introduced into
the cells 12 hours after sealing, only about 10% is reproduced.
Garrido and Rosenkranz (2004), therefore, the assumption
that an odor emitted by the larvae in the fifth stage is used as
signals for the mites to activate their ovaries. This chemical
benefits the receiver, cages were designed Special to confine
mites on various test objects, and found that mites trigger
oogenesis after perceiving pheromones emitted by larvae and
mites were deprived of food, as any bee hemolymph may also
contain signals. Extracts pentane cuticle of larvae also caused
activation of the ovaries of varroa, suggesting that the chemical
signal is polar. chemistry remains unidentified. Initially there
was a hypothesis that juvenile hormone in larvae / pupae bees
could be the factor which activates the ovaries of the mites, and
thus regulates reproduction. This

is an important hormone and

most insects that regulates oogenesis and spermatogenesis. This

theory was abandoned after observing differences in Africanized
bees and larvae of European honeybee, although it has been
shown that Africanized bees have reproductive rates much lower
mites (mainly due to a much higher level of infertile mites
percentage). When more than a mite invade a single cell
breeding, fertility decreases as the number of mites mother cell
increases. Mites invading brood cells in the combs of age but
also have fewer offspring. This led scientists to speculate that
the mites themselves may have a chemical that inhibits the
replication of each (a pheromone). A chemical ( Z)-8heptadecene, was identified. In the laboratory, there was a
30% reduction in mite fertility. When tested in the hive, the
average number of offspring was 3.48 in cells treated with (Z
)-8-heptadecene, but 3.96 in control cells. This difference was
small but statistically highly significant (P <0.01). Effective
fecundity (number of daughters potentially paired) was 0.94 in
cells treated, and 1.31 in the control cells, and this level of
difference should have a large impact on population growth to





start playback, many complex physiological processes must be in
place. Find critical genes for these processes can potentially
lead to new ways to control mites. Our laboratory recently
initiated a project to discover genes important for survival and
reproduction of mites, through the use of RNA interference
(RNAi). RNAi is a method to inject a relatively large
portion of dsRNA (400-500 bp long), which is cut into 20 to
30 bases long, then joins some complexes that eventually finds
complementary stretches of RNA and degrade them, resulting
in the reduction of mRNA of a target gene, ultimately its
protein product. Our basic principle is the pursuit of the same
genes that regulate the survival or reproduction in related
organisms (eg, ticks ) in the genome of the mite, synthesize
double-stranded RNA, inject RNA in mites, and then observe
their survival. If the injected mites survived, then proceeds to
observe its introducing breeding brood cells freshly capped.
Once a list of genes are found, then we need to ensure that the
dsRNA are specific to mites, and will not affect the bees, and
then find a way to introduce the dsRNA mites (either directly
or with the hemolymph of bees, which then pass the mites to feed
on the haemolymph of bees).

In summary, many factors can affect mite reproduction. These
range from the host reproductive rate (drone, worker or queen),
the cell size, the age of the larvae, the phoretic phase, relative
humidity, or even the movement of the combs. The more we
understand about how the reproduction of mites, the easier it
will be for us regulating find a way to interrupt the
reproduction of mites without harming the bees. The trick is
that the method must be easy and economical. Therefore, the
"basic research" on the reproductive biology of the mites will
eventually become useful for beekeepers, as it may one day
provide a new method for mite control.
Zachary Huang,

Department of Entomology, Michigan State University,





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