Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion
Some ethological aspects of A. mellifera Linnaeus, 1758 (Hymenoptera, Apidae), adaptation to parasiting the mite V. destructor Anderson et Trueman (Mesostigmata, Varroidae) are shown. The basic complexes of behaviour reactions, directed on a fight against the parasitic mites of bees brood at the gen...
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Інститут зоології ім. І.І. Шмальгаузена НАН України
2010
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| Цитувати: | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion / I.A. Akimov, V.E. Kiryushyn // Вестник зоологии. — 2010. — Т. 44, № 1. — С. 49–54. — Бібліогр.: 26 назв. — англ. |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1859730556274606080 |
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| author | Akimov, I.A. Kiryushyn, V.E. |
| author_facet | Akimov, I.A. Kiryushyn, V.E. |
| citation_txt | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion / I.A. Akimov, V.E. Kiryushyn // Вестник зоологии. — 2010. — Т. 44, № 1. — С. 49–54. — Бібліогр.: 26 назв. — англ. |
| collection | DSpace DC |
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| description | Some ethological aspects of A. mellifera Linnaeus, 1758 (Hymenoptera, Apidae), adaptation to parasiting the mite V. destructor Anderson et Trueman (Mesostigmata, Varroidae) are shown. The basic complexes of behaviour reactions, directed on a fight against the parasitic mites of bees brood at the genus Apis are shown, their comparative efficiency under various conditions and evolutional perspective. Possibility of ethological adaptation of honey bee to V. destructor parasiting, direction of selection by this sign and influencing of human on parasitic-host system was discussed. An approach to the selection of bees with the purpose of resistanse to varroosis promoution is proposed.
В статье освещаются некоторые этологические аспекты адаптации медоносной пчелы Apis mellifera Linnaeus, 1758 (Hymenoptera, Apidae) к паразитированию на ней клеща Varroa destructor Anderson et Trueman (Mesostigmata, Varroidae). Показаны основные комплексы поведенческих реакций, направленных на борьбу с клещами-паразитами расплода у пчел рода Apis, их сравнительная эффективность в различных условиях и эволюционная перспективность. Обсуждается возможность этологической адаптации медоносной пчелы к варроа, направленность отбора по этому признаку и влияние человека на данную паразито-хозяинную систему. Предложен подход к селекции пчел с целью повышения устойчивости к варроозу.
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UDC 595.42:595.799
ETOLOGICAL ASPECTS OF HONEYBEE APIS MELLIFERA
(HYMENOPTERA, APIDAE) ADAPTATION TO PARASITIC
MITE VARROA DESTRUCTOR (MESOSTIGMATA,
VARROIDAE) INVASION
I. A. Akimov, V. E. Kiryushyn
Shmalhausen Institute of Zoology NAS of Ukraine
B. Chmielnicky str., 15, Kyiv, 01601 Ukraine
E-mail: kyrya1@yandex.ru
Received 3 November 2009
Accepted 2 December 2009
Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite
Varroa destructor (Mesostigmata, Varroidae) Invasion. Akimov I. A., Kiryushyn V. E. — Some ethological
aspects of A. mellifera Linnaeus, 1758 (Hymenoptera, Apidae), adaptation to parasiting the mite
V. destructor Anderson et Trueman (Mesostigmata, Varroidae) are shown. The basic complexes of
behaviour reactions, directed on a fight against the parasitic mites of bees brood at the genus Apis are
shown, their comparative efficiency under various conditions and evolutional perspective. Possibility of
ethological adaptation of honey bee to V. destructor parasiting, direction of selection by this sign and
influencing of human on parasitic-host system was discussed. An approach to the selection of bees with
the purpose of resistanse to varroosis promoution is proposed.
K e y wo r d s: Apis mellifera, Varroa destructor, hygienic behaviour of bees, resistance, adaptation,
selection.
Ýòîëîãè÷åñêèå àñïåêòû àäàïòàöèè ìåäîíîñíîé ï÷åëû, Apis mellifera (Hymenoptera, Apidae),
ê ïàðàçèòèðîâàíèþ íà íåé êëåùà Varroa destructor (Mesostigmata, Varroidae). Àêèìîâ È. À.,
Êèðþøèí Â. Å. —  ñòàòüå îñâåùàþòñÿ íåêîòîðûå ýòîëîãè÷åñêèå àñïåêòû àäàïòàöèè ìåäîíîñíîé
ï÷åëû Apis mellifera Linnaeus, 1758 (Hymenoptera, Apidae) ê ïàðàçèòèðîâàíèþ íà íåé êëåùà
Varroa destructor Anderson et Trueman (Mesostigmata, Varroidae). Ïîêàçàíû îñíîâíûå êîìïëåêñû
ïîâåäåí÷åñêèõ ðåàêöèé, íàïðàâëåííûõ íà áîðüáó ñ êëåùàìè-ïàðàçèòàìè ðàñïëîäà ó ï÷åë ðîäà
Apis, èõ ñðàâíèòåëüíàÿ ýôôåêòèâíîñòü â ðàçëè÷íûõ óñëîâèÿõ è ýâîëþöèîííàÿ ïåðñïåêòèâíîñòü.
Îáñóæäàåòñÿ âîçìîæíîñòü ýòîëîãè÷åñêîé àäàïòàöèè ìåäîíîñíîé ï÷åëû ê âàððîà, íàïðàâëåííîñòü
îòáîðà ïî ýòîìó ïðèçíàêó è âëèÿíèå ÷åëîâåêà íà äàííóþ ïàðàçèòî-õîçÿèííóþ ñèñòåìó. Ïðåäëîæåí
ïîäõîä ê ñåëåêöèè ï÷åë ñ öåëüþ ïîâûøåíèÿ óñòîé÷èâîñòè ê âàððîîçó.
Êëþ÷åâûå ñ ëîâ à: Apis mellifera, Varroa destructor, ãèãèåíè÷åñêîå ïîâåäåíèå ï÷åë, àäàïòàöèÿ,
ñåëåêöèÿ.
Introduction
It is impossible to study honeybee adaptation paths to Varroa destructor Anderson et Trueman, 2000
parasitism without examination of the mite and bee population interactions as a whole. It should be done because
of host-parasite relationships in mite — bee system being not fully fixed and are being formed nowadays due
to recent switching of V. destructor to parasitizing honeybee (Akimov et al., 1993). Previously, species from
the genus Varroa parasitized other species from the genus Apis including Apis cerana F. At the same time, it is
known that Asiatic and European honey bees are quite similar biologically and they readily get into their colonies
bees from the other species brood. This fact and direct observations indicate that behavioral acts and their
complexes, as well as communication and information transmission systems among individuals in colonies of
these species are similar in many aspects (Rath, 1993; Kuznetsov, 2005).
Being similar in general, European and Asian honey bees have different thresholds and levels of
ethological reactions common to both species, and these appear to allow Asian honey bee to control parasite
population. For example, Asian honey bee has greater excitability and propensity to slough-offs and, in
Vestnik zoologii, 44(1): 49–54, 2010
addition, it has less diameter of comb cells and slightly shorter period of brood development (Peng et al., 1987;
Kuznetsov, 2005). However, both species have inherent reactions fn body cleaning as a response to irritation,
and are also able to identify cells with mity brood. All this determines the Asian honey bee adaptive reactions
complexity to Varroa parasitism and causes certain methodical complications in studying them as compared
to those reactions in European honeybee.
As possible preadaptations, one should consider different growth rate of parasite number in colonies of
different European honeybee breeds and populations, which may depend on different size of honeycomb cells
(Harbo, Harris, 1999; Rosenkranz, 1999) or some asynchronies in development of mite and pupae in brood
(Akimov et al., 1993).
Results
Complex studies of the parasite biology and different aspects of its interactions with
European honeybee allow focusing mainly on such kinds of bees resistance to the para-
site as their adaptive behavioral reactions studied by us and other authors. (Akimov et
al., 1993; Akimov, Kiryushyn, 2008; Ritter et al., 1990; Rosenkranz et al., 1997; Spivak,
Gilliam, 1998; Correa-Marques, de Jong, 1998; Moretto, Melco, 2001; Fries et al., 2006;
Fries, Bommarco, 2007).
Such reactions consist of three basic behavioral complexes. The first of these behav-
ioral complexes is the most primitive, mostly observed in Asian honey bee. It consists in
slough-off of all bee imaginal forms. At that combs and brood are abandoned together
with parasites, most part of which remains in the brood. Such extraordinary response takes
place when there is abundant number of parasites considerably disturbing bees (Kuznetsov,
Proschalycin, 2004; Kuznetsov, 2005).
Slough-offs at varroatosis is described for European honeybee as well (Rosenkranz,
1999; Khmara, 2002). Primitiveness of such reaction consists in heavy losses. Bees have
to completely restore the nest and accumulate new feedstuffs. Such technique of colony
cleaning is successful only if long-term honey gathering in nature is possible, which enables
accumulation of feedstuffs for winter and rearing enough number of bees. In fact, bee
slough-offs are possible during swarming only, however, the extensiveness of bees infec-
tion with the parasite is relatively low at this period. The number of mites on adult bees
increases in August and September when forage gathering appears to be impossible. The
fact of autumn slough-offs is an indirect evidence of bee tropical origin, as it is in the
South Asia where the stable honey gathering in October and November is observed, and
where the vast majority of species from the genus Apis is described. Such kind of honey
gathering is not observed in the Eastern Europe, where, as a rule, it is finished in
August — beginning of September. In the temperate climate of the Eastern Europe, the
autumn slough-offs may result in bees death during winter, so such form of adaptation
to varroa can not be prospective, and bee population under our conditions will selective-
ly be cleaned from colonies tending to slough-offs.
The second complex of behavioral reactions associated with adaptations to varroa par-
asitizing is the development of bee body cleaning reactions investigated by us and other
authors. Such behavioral reactions in bees are divided into self-cleaning and mutual clean-
ing, similar to grooming in higher vertebrates, when one bee cleans another one (Akimov,
Kiryushyn, 2008). Self-cleaning of adult Asian honey bees aimed at relieving from var-
roa mites was described by Y. S. Peng (Peng et al., 1987). It is a modification of the typ-
ical body cleaning movements and is used when the mite is present on bee’s body (Rath,
1993). It is necessary to note that the body cleaning movements are rather conservative
and only slightly in different species not only in Apis, but also in Apoidea on the whole.
Cleaning of body vestiture is a typical reaction of any insect on body contamination,
mechanical irritation or stress. Even with the increasing of insects sociality, such reac-
tions remain almost unchanged (Michener, 1974).
On the other side, the reactions of body self-cleaning observed in European hon-
eybees are not specific. The complex of movements starts at the presence of any irritant.
50 I. A. Akimov, V. E. Kiryushyn
A bee gets rid from a mite only in insignificant number of cases (Kiryushyn, 2005 a).
Highly conservative movements of body vestiture cleaning suppose that the Asioan
honey bee have the same pattern as seen in the other bees. Probably, their threshold val-
ues of irritation are lower than in the European honeybee, so they do more acts of self-
cleaning in mite presence and chances to throw the mite off or to damage it also
increase.
It is known, that with the insect sociality increase, the division of labor in a nest
increases as well (Levchenko, 1976). Since the European honeybee have the largest colonies
with the greatest degree of labor division and information exchange between colony mem-
bers as compared to other bees, we may logically assume new behavioral models at irri-
tation, e. g. mutual body cleaning, built-in the system of age and caste labor division in
a bee nest, and specific, directed against varroa, self-cleaning techniques.
The comprehensive study of honeybee body cleaning reactions found out their
dependence of the environment temperature, honey gathering, and stress factors. Thus,
the highest intensity of such reactions was observed at temperatures close to optimum
for bee colony: 30–35°Ñ. Consequently, honey gathering is increased by the bees clean-
ing their bodies, however its intensity in some individuals reduces. We confirmed the pres-
ence of developed reactions of both body self-cleaning and mutual cleaning in bees. Also,
within this study, we established the role of age-dependent bee polyethism in develop-
ment of such reactions (Kiryushyn, 2005 b). Under significant stress irritations, such as
mechanical effect or bee “narcotization”, the reactions of body cleaning in bees were con-
siderably activated. However, such reactions of body cleaning appeared to be nonspecif-
ic and ineffective against varroa. Moreover, within an hour after the parasite introduc-
tion, bees stopped cleaning movements that may be explained by the CNS inhibition devel-
oped during a long-term irritation. This significantly reduced reactions efficiency, as the
intensity of movements rose only for a short time after the mite transfer to another bee,
but went down quickly.
Thus, we may speak about potentially possible European honeybee adaptation to the
varroa infection by the development of body cleaning reactions such as those seen in Asian
honey bee, however, at this stage, such behaviour is not able to restrain parasite repro-
duction due to its unspecificity, low efficiency and rapid extinguishing. At the same time,
the parasite easily recognizes the main host, that makes possible its transfer among bees
of the same colony. It also gives the mite an opportunity to infect bees and remain on
them up to ceasing of the body cleaning reaction.
As far as the primary reactions already exist, their adaptive specialization against var-
roa irritation may evolve within an evolutionary insignificant period of time. According
to our data, the presence of varroa on an adult bee induces, at this stage, general acti-
vation of body cleaning reactions, which, however, are not specific and, being the same
as the reactions against mechanical or stress irritation, remain ineffective.
The third complex of behavioral reactions against varroa is the bee colony particu-
lar control of varroa population in brood. Since the varroa reproduction and develop-
ment occur in brood, the most effective control method for the bee colony could be recog-
nition and elimination of the parasites during their development in the brood. It would
result in the lowest loss rates for colony as compared to other control methods. Moreover,
larval stages of mite in beehive environment, out of brood cells, are unviable, therefore
their elimination would not require bees considerable efforts.
Our own data and literary information show that bees have certain mechanisms for
recognition of affected brood, which, however, are studied insufficiently (Rosenkranz et
al., 1997; Spivak, Gilliam, 1998). Such differences may be due to the development of
secondary microflora on pupae, weakened by the varroa parasitism. On the other hand,
the connection between the hygienic capabilities of bee colonies and rates of growing of
varroa population may be mediated (Maslennikova, 2003).
51Ethological Aspects of Honeybee Apis mellifera…
Brazilian bees and some lines of Italian bees partly throw out pupae affected by var-
roa, however, it is not effective mechanism to control the parasite population and when
extensiveness of brood affection is over 5% such reaction goes down or stops complete-
ly (Rosenkranz et al., 1997).
Recognition and the following unsealing and rapid elimination of the affected and
dead larvae and pupae from cells is one of the main methods to control brood diseases
among bees. Mechanisms of such recognition are not clear enough, however its great effi-
ciency against many diseases are doubtless (Rudenko, 2004; Spivak, Gilliam, 1998).
We found out great variability of this trate in a group of bee colonies.
During the study of growth rates of varroa population in some bee colonies we found
out the dependence between extensiveness of individual infection and hygienic capabil-
ities of bee colonies. Namely, colonies, where bees recognized cells with the infected brood
in a shorter period of time, had lower growth rate of varroa population during three bee-
keeping seasons and had significantly lower extensiveness of adult bees infection after leav-
ing the brood cells (Kiryushyn, 2007 a; Akimov, Kiryushyn, 2008).
However, even in the colonies with the greatest hygienic capabilities the number of
mites grows so quickly that may result in death of the colony in two seasons after the
last treatment with acaricide. Some parameters of winter resistance in bee colonies cor-
relate with their hygienic capabilities. So, when analyzing winter resistance of bee
colonies with the different level of hygienic capabilities, some differences were observed
such as colony attenuation in wintering period and forage consumption of bees. Moreover,
there is a correlation between the level of hygienic capabilities and brood amount in bee
colonies after flying around, probably due to different way of colony development in the
period before flying around (Kiryushyn, 2007 b).
Thus, as in the case of body cleaning reactions, bee colonies have already had mech-
anisms for certain control of varroa population in brood, however they are not specific
to the mite and just slow down increasing of the parasite population in a colony. On the
other hand, the bee colonies able to rapidly recognize infected brood and maintain accept-
able sanitary condition of nests, in addition to the lower growth rate of V. destructor pop-
ulation inside them also have a complex of other trates useful for surviving, such as high-
er winter resistance, earlier colony activation in spring, greater resistance to other dis-
eases (Kiryushyn, 2008; Rudenko et al., 2006).
As is obvious from the forementioned, bee adaptations to varroa parasitism are with-
in adaptations to many unfavorable environmental factors and are only slightly modified
depending on these factors. Therefore hygienic activity, winter resistance, and other reac-
tions towards decreasing of parasite reproduction rates and its effect on the bee colony
became vital. It is especially important to emphasize, that natural resistance of Asian honey
bee known to be resistant to varroosis is initially limited by biological characteristics of
European honeybees and their living conditions.
Our data gives us certain hope on gradual spontaneous increase in bee resistance to
varroa invasion. Moreover, bees have principle possibility to slow down parasite repro-
duction by decreasing cell diameter to 4.7–4.9 mm, and reducing period of pupa devel-
opment, or changing the timing of its hormonal transformations. However, it may
appear to be another biologically or economically limiting factor. Anyway, at this stage
we can not see anything of this kind even at hands of man (Maslennikova, 2003;
Kulincevuc, 1987; Moretto, 2001).
Honeybee adaptation to varroa parasitizing seems to still be in on the first phases
of emerging, however there are approximate paths and specific mechanisms of subsequent
mutual bee and mite adaptation to the stable coexistence, as well as man’s role in this
process. It is also obvious that the control on varroa population in a bee nest develops
as a difficult complex of different and disconnected trates inherited with participation of
the large number of genes. Therefore, bee selection to varroa resistance is confronted with
52 I. A. Akimov, V. E. Kiryushyn
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54 I. A. Akimov, V. E. Kiryushyn
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| id | nasplib_isofts_kiev_ua-123456789-65659 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0084-5604 |
| language | English |
| last_indexed | 2025-12-01T12:48:14Z |
| publishDate | 2010 |
| publisher | Інститут зоології ім. І.І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Akimov, I.A. Kiryushyn, V.E. 2014-06-30T07:40:06Z 2014-06-30T07:40:06Z 2010 Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion / I.A. Akimov, V.E. Kiryushyn // Вестник зоологии. — 2010. — Т. 44, № 1. — С. 49–54. — Бібліогр.: 26 назв. — англ. 0084-5604 https://nasplib.isofts.kiev.ua/handle/123456789/65659 595.42:595.799 Some ethological aspects of A. mellifera Linnaeus, 1758 (Hymenoptera, Apidae), adaptation to parasiting the mite V. destructor Anderson et Trueman (Mesostigmata, Varroidae) are shown. The basic complexes of behaviour reactions, directed on a fight against the parasitic mites of bees brood at the genus Apis are shown, their comparative efficiency under various conditions and evolutional perspective. Possibility of ethological adaptation of honey bee to V. destructor parasiting, direction of selection by this sign and influencing of human on parasitic-host system was discussed. An approach to the selection of bees with the purpose of resistanse to varroosis promoution is proposed. В статье освещаются некоторые этологические аспекты адаптации медоносной пчелы Apis mellifera Linnaeus, 1758 (Hymenoptera, Apidae) к паразитированию на ней клеща Varroa destructor Anderson et Trueman (Mesostigmata, Varroidae). Показаны основные комплексы поведенческих реакций, направленных на борьбу с клещами-паразитами расплода у пчел рода Apis, их сравнительная эффективность в различных условиях и эволюционная перспективность. Обсуждается возможность этологической адаптации медоносной пчелы к варроа, направленность отбора по этому признаку и влияние человека на данную паразито-хозяинную систему. Предложен подход к селекции пчел с целью повышения устойчивости к варроозу. en Інститут зоології ім. І.І. Шмальгаузена НАН України Вестник зоологии Экология Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion Этологические аспекты адаптации медоносной пчелы, Apis mellifera (Hymenoptera, Apidae), к паразитированию на ней клеща Varroa destructor (Mesostigmata, Varroidae) Article published earlier |
| spellingShingle | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion Akimov, I.A. Kiryushyn, V.E. Экология |
| title | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion |
| title_alt | Этологические аспекты адаптации медоносной пчелы, Apis mellifera (Hymenoptera, Apidae), к паразитированию на ней клеща Varroa destructor (Mesostigmata, Varroidae) |
| title_full | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion |
| title_fullStr | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion |
| title_full_unstemmed | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion |
| title_short | Ethological Aspects of Honeybee Apis mellifera (Hymenoptera, Apidae), Adaptation to Parasitic Mite Varroa destructor (Mesostigmata, Varroidae) Invasion |
| title_sort | ethological aspects of honeybee apis mellifera (hymenoptera, apidae), adaptation to parasitic mite varroa destructor (mesostigmata, varroidae) invasion |
| topic | Экология |
| topic_facet | Экология |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/65659 |
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