COLONY (Apis mellifera L.)
Z b i g n i e w
L i p i ñ s k i
10-736 Olsztyn, Janina Wengris str 8. Poland. E-mail: [email protected] Received 05 April 2007; accepted 08 May 2007 The observation that withdrawal of the queen from the colony of western honeybees (A. mellifera L.) cause it unusual agitation connected with subsequent ovary activation and un-fertilized egg laying by sterile workers (W i n s t o n 1987) indicates that the calming effect ofqueen substance (understood as a full blend of its pheromones) enforced by a similarly actingblend of open brood pheromones known as “pheromonal queen control”, is the main device ofreproductive dominance of the queen. Consider that emotional system of the honeybee governsand orchestrates it behaviour with physiology.
Based on this hypothesis the author presents possible signaling pathways of this reproductory dominance in context of so called Lee-Booth effect. Last findigs indicate that worker bee tissuescontain some “higher” animal reproductory (FSH, LH) and sex hormones (testosterone, E2). Inview of above hypothesis the author concludes that role of evolutionary conserved gonadotropicand sex hormones in regulation of reproduction in honeybees should be the object of more in-tensive studies.
Keywords: honey bee, honeybee queen, reproduction, FSH, LH, GnRH, E 2, Lee-Boot
are normally sterile, but under certain con- neuronal circuits of the emotional system ditions this is not true (Bourke 1988). For instance the honeybee workers Apis mellifera L. usually do not reproduce, but sider that emotional system of the honey- can activate their ovaries under queenless bee can govern and orchestrate it behaviour rule, laying workers of the Cape honeybee, emotional piping of queenless laying work- A. m. capensis Esch., can parthenogetically ers of A. m. ligustica (O h t a n i and action of surplus of emotional agitation.
bees (A. mellifera L.) cause it unusual agi- PATHWAYS OF SUPPRESSION OF
tation (reviewed by L i p i n s k i 2006 a, WORKER REPRODUCTION
2001) connected with ovary activation and unfertilized egg laying by sterile workers (QS) – understood as a full blend of her (Wi n s t o n 1987) indicates, that repro- ceptive abilities of workers (reviewed by L i p i ñ s k i 2006 a). Especially in stress of innate behaviours (instincts) (Lipiñski conditions (Lin et al. 2004). For instance 2006 a) (3) – NCES orchestrate these be- its high level enhance perceptive abilities L i p i ñ s k i 2006 a), an assumption arise ing/emotional reason, in queenless colonies building, queen rearing etc.) and physiol- of less stress sensitive European workers ogy of ovary activation and egg laying as ovary development did not begin until most stress sensitive Africanized workers (more resistant to this calming effect) while un- field (G r o z i n g e r et al. 2003) and by ley et al. 2003) indicate that in some cases this suppression is not strong enough.
tion of bees which makes them more resis- tant to this calming effect (L i p i ñ s k i 2001). This explanation confirms fact that al. 1992) in queenless colonies have a low sQMP do not inhibit quen rearing in terms level of JH-3 titetrs in their haemolymph, (2) - JH-3 do not increase ovary develop- Lipiñski 2001) and do not suppress ovary development in (highly agitated) queenless (R o b i n s o n et al. 1992), (3) – JH-3 colonies of A. mellifera L., (Willis et al.
vitellogenin synthesis (L u c i m a r a et al.
2000), (4) - JH-3 do not make ovary activa- cussed calming suppression of workers re- tion and egg laying (Mohameddi et al.
production confirms also, that: not only a 1998, W i n s t o n and S l e s s o r 1998) natural blend of queen pheromones (QS) – strongly suggest that there is no evidence that JH-3 acts as a traditional gonadotropin honeybee JH-3 has partly lost in evolution brood (BP) can suppress, ovary activation its gonadotropic effect if it ever was.
L i p i ñ s k i 2006 b) and elevate their re- sponse thresholds (P a n k i v et al. 1998).
Significantly for this calming phenomenon, both QS (Kaatz et al. 1992, Pankiv et al.
cluded, because low JH-3 titers in workers suppress juvenile hormone (JH-3) and slow vitellogenin (Pinto et al. 2000). The more so that, reproductivelly dominant individu- is because JH-3, as mediator of emotional als of A. m. capensis, which are resistant to responses of honeybees can influence per- poorly understood (G r i m m e l i k h u i j z e n et al. 2000). Control of gonad development ecdysteroids and a peptidic brain gonado- tropins (D e L o o f et al. 2001). What is reproductory) dominance of the queen over tion that emotional piping of workers after queeen loss (Ohtani and Kamada 1980).
insects with respect to the molecular struc- (Sasaki and Nagao 2002) in brain levels.
ture of gonadotropins is far less uniform.
ovary activation in queenless Apis mellifera are glycoproteins that are synthezised and biosynthesis (Robinson et al. 1992). It is noteworthy that correlation between brain tion of progesterone or of sex steroids (De intercerebralis of the insect brain regulates Interestingly it is as result of that of ex- certain stages of the reproductive process usual behaviours appear. For instance: (1) - more in Drosophila melanogaster, two irregular egg laying, (2) - emergency queen G-protein coupled receptors, structurally cells construction with dead drone larvae related to the mammalian glycoprotein hor- (3) - small cups built over pollen or (4) - an egg is laid on the pollen mass (Morse and vertebrate nervous systems appear to utilize mechanism of calming suppression of work- similar intercellular signal molecules” ers reproduction discussed above confirms (Stefano 2002), (2) – glycoproteine go- also fact that signals arise from the tergal glands of queens can also inhibit worker re- production (Wossler and Crewe 1999).
2002), (3) – measurable immunore-activi- THE POSSIBLE ROLE OF
ties characteristic for sex hormones such as GLIKOPROTEINE HORMONES IN
testosterone (T) and estradiol-17 beta (E 2) REGULATION OF HONEYBEE
– testosterone was found in royal jelly started relatively late (C y m b o r o w s k i production in mature female of Bombyx nisms behind reproduction are, so far, only mori (Ohnishi et al. 1985), an assumptionarises that “unsolved problem of hormonal Neobellieria bullata (T h u e n i s et al.
1989), (5) - levels of LH, and E 2 in worker bees emerged and developed in the absence can be solved base on hypothesis that this of the queen, rose at the third day (Zhou and Wen 2002), (5) – the content of FSH of nursing bees from the day of emergence LH, E 2, testosterone or even gonadotropin is more or less the same with a decreasing trend, while it is significantly higher in for- 500 milion years of evolution (Chen et al.
H a u s e r et al. (2000) who cloned from Drosophila melanogaster receptor that is structurally and evolutionary related to the found that the frut fly produces at least two ancestor with humans was 600 milion years evolutionarily related to the LH and FSH. It peats containing G protein-coupled recep- changes in social context (R o b i n s o n tor-1 which transduce extracellular signals into cellular physiological responses) and these receptors are involved in the control last two hormones directly regulate ovary of insect reproduction (D e L o o f et al.
other similar releasing hormone) by QP and Wo o d r i n g 1992, B i c k e r et al. 1988) as well. Especially because: (1) - by using monoclonal antibodies directed against dif- receptors resulting in the influx of Ca 2+, ferent epitopes of human LH, Ve r h a e r t (NOS) activity, in a situation when nitric americana, (3) De Loof et al. (2001) ob- et al. 1997). Consider that glutamate is be- brain extract on testosterone production by mouse Leyding cells (4) – LH and FSH – THE POSSIBLE ROLE OF THE
ies and deliberations I venture to put for- ward the hypothesis that the calming effectof QS, understood as a full blend of its pheromones enforced by a similarly acting the regulation of reproductive development often involves chemical signals that either advance or delay maturation (S t e r n and “pheromonal queen control” (Keller and gen-dependent chemical activators acceler- and behavioural dominance of the queen in gland-dependent inhibitors delay it. These signals are thought to be perceived by the oprojections to the accessory olfactory bulb.
duction in honeybees should be the object higher brain centres that ultimately controlthe release of ovarian luteinizing hormone.
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Z b i g n i e w
L i p i ñ s k i
Dobrze znany fakt, ¿e usuniêcie matki z rodziny pszczelej powoduje nadzwyczajne pobu- dzenie nerwowe robotnic wraz z póŸniejszym sk³adaniem przez nie niezap³odnionych jaj(W i n s t o n 1987) wskazuje, ¿e uspokajaj¹cy wp³yw feromonów matki, wspomagany przezpodobnie dzia³aj¹ce feromony m³odego czerwiu, znany jako „feromonalna kontrola ze stronymatki” (ang. pheromonal queen control), stanowi g³ówny mechanizm reprodukcyjnej dominacjimatki w rodzinie pszczelej. Maj¹c jednoczeœnie na uwadze, ¿e system emocjonalny pszczo³ymiodnej steruje jej zachowaniami, zgrywaj¹c je z odpowiednimi reakcjami fizjologicznymi.
Bazuj¹c na tej hipotezie, autor przedstawia mo¿liwe drogi jakimi substancje sygnalne steruj¹ t¹ dominacj¹ w kontekœcie efektu Lee-Booth’a. Potwierdzeniem tej hipotezy wydaj¹ siê byæostatnie odkrycia wskazuj¹ce, ¿e tkanki pszczo³y miodnej zawieraj¹ typowe dla tzw. „zwierz¹twy¿szych” hormony gonadotropowe (FSH, LH) oraz p³ciowe (estradiol - E 2 testosteron).
W œwietle powy¿szej hipotezy autor wnioskuje, ze rola ewolucyjnie starych hormonów gonadotropowych i p³ciowych w regulacji rozmna¿ania siê pszczó³ winna staæ siê obiektem
wzmo¿onych badañ
S³owa kluczowe: pszczo³a miodna, matka pszczela, FSH, LH, GnRH, E 2, Efekt

Source: http://www.jas.org.pl/jas_51_1_2007_11.pdf


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