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1.
Worker-reproduction is rare in queenright honey-bee colonies. When workers do lay eggs, their eggs are normally eaten by other workers presumably because they lack the queen's egg-marking signal. Workers use the absence of this queen signal to enforce the queen's reproductive monopoly by policing any worker-laid eggs. In contrast, in anarchistic colonies, the majority of the males arise from worker-laid eggs. Anarchistic worker-laid eggs escape policing because workers perceive anarchistic eggs as queen-laid. However, in this study, we show that eggs laid by queenless anarchistic workers do not escape policing and have very similar removal rates to worker-laid eggs from queenless wild-type (i.e. non-anarchistic) colonies. This suggests that, under queenless conditions, eggs laid by anarchistic workers lose their chemical protection and are therefore no longer perceived as queen-laid. Hence, the egg-marking signal seems to be only applied to eggs when queen and brood are present. This suggests that in the absence of queen and brood, the biosynthetic pathway that produces the egg-marking signal is switched off.Communicated by L. Keller 相似文献
2.
Informational constraints can be an important limitation on the accuracy of recognition. One potential constraint is the use
of recognition information from the same sources in multiple discriminatory contexts. Worker wood ants, Formica fusca, discriminate eggs based on their maternal sources of origin in two main contexts: recognition of eggs laid by nestmate versus
non-nestmate queens and recognition of worker-laid versus queen-laid eggs. We manipulated the experience of F. fusca workers in laboratory colonies to both worker-laid and queen-laid eggs by transferring eggs between colonies in order to
investigate whether these two contexts of egg discrimination are independent. Experience of non-nestmate queen-laid eggs significantly
increased worker acceptance of both familiar (18% accepted) and unfamiliar (10%) queen-laid eggs compared to control workers
without experience of eggs other than those laid by their own colony’s queen (2%). In contrast, worker acceptance of worker-laid
eggs was not affected by variation in the egg experience of workers (14% in workers from control colonies exposed only to
eggs from their own colony’s queen versus 19% and 17% in workers from colonies which had received eggs laid by either a non-nestmate
queen or nestmate workers, respectively). Our results suggest that these two recognition contexts do not strongly constrain
each other and are different in their ontogeny. In particular, worker-laid eggs are universally discriminated against by workers
from colonies with a queen whatever the egg experience of the workers, while non-nestmate queen-laid eggs are strongly discriminated
against only by workers without experience of eggs laid by more than one queen. 相似文献
3.
Benjamin P. Oldroyd Luke A. Halling Gregory Good Wandee Wattanachaiyingcharoen Andrew B. Barron Piyamas Nanork Siriwat Wongsiri Francis L. Ratnieks 《Behavioral ecology and sociobiology》2001,50(4):371-377
Workers of the Asian hive bee, Apis cerana, are shown to have relatively high rates of worker ovary activation. In colonies with an active queen and brood nest, 1-5% of workers have eggs in their ovarioles. When A. cerana colonies are dequeened, workers rapidly activate their ovaries. After 4 days 15% have activated ovaries and after 6 days, 40%. A cerana police worker-laid eggs in the same way that A. florea and A. mellifera do, but are perhaps slightly more tolerant of worker-laid eggs than the other species. Nevertheless, no worker's sons were detected in a sample of 652 pupal males sampled from 4 queenright colonies. A cerana continue to police worker-laid eggs, even after worker oviposition has commenced in a queenless colony. 相似文献
4.
In anarchistic honey-bee colonies, many workers’ sons are reared despite the presence of the queen. Worker-laid eggs are normally
eaten by other workers in queenright colonies. Workers are thought to discriminate between queen-laid and worker-laid eggs
by the presence or absence of a queen-produced egg-marking pheromone. This study compared the survival of three classes of
eggs (worker-laid eggs from anarchistic colonies, worker-laid eggs from non-anarchistic queenless colonies, and queen-laid
eggs) in both queenright normal colonies and queenright anarchistic colonies, in order to test the hypothesis that anarchistic
workers evade policing by laying more acceptable eggs. As expected, few worker-laid eggs from non-anarchistic colonies survived
more than 2 h. In contrast, worker-laid eggs from anarchistic colonies had much greater acceptability, which in some trials
equalled the acceptability of queen-laid eggs. Anarchistic colonies were generally less discriminatory than normal queenright
colonies towards worker-laid eggs, whether these originated from anarchistic colonies or normal queenless colonies. This indicates
that the egg-removal aspect of the anarchistic syndrome involves both worker laying of eggs with greater acceptability and
reduced discriminatory behaviour of policing workers.
Received: 19 July 1999 / Received in revised form: 3 November 1999 / Accepted: 20 November 1999 相似文献
5.
Nestmate recognition for eggs in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)
Christian W. W. Pirk Peter Neumann Randall Hepburn 《Behavioral ecology and sociobiology》2007,61(11):1685-1693
Colony integrity is fundamental to social insects and is threatened by the reproduction of non-nestmates. Therefore, discrimination
between eggs derived from nestmates and non-nestmates would constitute an adaptation to prevent exploitation of the entire
cooperative group by unrelated individuals. The removal of nestmate and non-nestmate queen and worker-laid eggs was evaluated
in honeybees using colonies of Apis mellifera capensis to test female and of A. m. scutellata to test male eggs. The data show that honeybees can distinguish between nestmate and non-nestmate eggs of both sexes. Moreover,
non-nestmate female queen-laid eggs were removed significantly faster than nestmate female worker-laid eggs in A. m. capensis, indicating that nestmate recognition cues can override caste-specific ones. While the experimental manipulation accounts
for 37.2% (A. m. scutellata) or 1.6% (A. m. capensis) of variance in relation to egg removal, nestmate recognition explains 33.3% for male eggs (A. m. scutellata) and 60.6% for female eggs (A. m. capensis), which is almost twice as high as the impact of caste (16.7% A. m. scutellata; 25% A. m. capensis). Our data show a stronger effect of nestmate recognition on egg removal in the honeybee, suggesting that cues other than
caste-specific ones (viability/kin) can dominate egg removal behavior. In light of intraspecific social parasitism, preventing
the reproduction of unrelated individuals (group selection) rather than preferring queens’ eggs (kin selection) appears to
be the driving force behind the evolution of egg removal behavior in honeybees. 相似文献
6.
Wim Bonckaert Adam Tofilski Fabio S. Nascimento Johan Billen Francis L. W. Ratnieks Tom Wenseleers 《Behavioral ecology and sociobiology》2011,65(4):633-640
In insect societies, workers often try to challenge the reproductive monopoly of the queen by laying their own eggs. Successful
worker reproduction, however, is frequently prevented by queen policing or worker policing, whereby either the mother queen
or non-reproductive workers selectively kill worker-laid eggs. Recently, a third mechanism—“selfish” worker policing—has also
been described in which the workers selectively police worker-laid eggs but also lay eggs themselves. Here, we present results
from the monogynous wasp Dolichovespula norwegica, which show that all three kinds of policing—queen policing, worker policing and “selfish” worker policing—co-occur. The
net effect of these three kinds of policing collectively favoured the queen’s reproduction, as within 1 day 44% of the worker-laid
eggs versus only 8% of the queen-laid eggs were eaten. Of the worker-laid eggs that were killed by workers, approximately
two thirds were eaten by the reproductive workers even though these made up only a small proportion, 8%, of the work force.
This means that policing workers obtained both direct fitness benefits as well as indirect (inclusive) fitness. In addition,
we show that worker policing was carried out by a limited, specialised set of workers that was estimated to constitute approximately
one quarter of the whole colony and of which 66% were non-reproductive. 相似文献
7.
Nathalie Stroeymeyt Elisabeth Brunner Jürgen Heinze 《Behavioral ecology and sociobiology》2007,61(9):1449-1457
Animal societies, including those of humans, are under constant threat by selfish individuals, who attempt to enforce their
own interests at the cost of the group. In the societies of bees, wasps, and ants, such individual selfishness can be prevented
by “policing,” whereby workers or queens impede the reproduction of other individuals by aggression, immobilization, or egg
eating. In this study, we report on a particular kind of reproduction control in the ant Temnothorax unifasciatus, which can be considered as a selfish act itself. We experimentally induced workers to lay eggs by dividing several colonies
into two halves, one with and one without a queen. In queenless colonies, workers established rank orders by aggression and
several top-ranking workers started to reproduce. Upon reunification, egg-laying workers mostly stopped behaving aggressively.
They were neither attacked by the queen nor by random workers, but instead received infrequent, nondestructive, targeted aggression
from a few workers, most of which became fertile when the queen was later removed. The introduction of differentially stained
worker-laid and queen-laid eggs in queenright fragments did not lead to a selective removal of worker-laid eggs. Hence, there
appears to be no collective worker policing in T. unifasciatus. Instead, reproduction appears to be controlled mostly through a few attacks from high-ranking workers, which, in this way,
might attempt to selfishly increase their chances of future reproduction. 相似文献
8.
Heikki Helanterä Stephen J. Martin Francis L. W. Ratnieks 《Behavioral ecology and sociobiology》2007,62(2):223-228
We studied the effect of prior experience to eggs laid by nestmate and non-nestmate queens on the acceptance of queen-laid
eggs by worker wood ants, Formica fusca. We transferred eggs from a non-nestmate queen into colonies during early spring, when their own queen was recommencing egg
laying. A few weeks later, workers from these “experienced” colonies accepted eggs of both familiar (44% acceptance) and unfamiliar
(40%) non-nestmate queens much more than workers from control colonies (2%) that had only had previous contact with their
own queen’s eggs. Thus, prior exposure to eggs laid by a non-nestmate queen induces much greater acceptance of all non-nestmate
queen-laid eggs. Mechanistically, we hypothesize that exposure to eggs from several queens may increase acceptance by causing
a highly permissive acceptance threshold of non-nestmate queen-laid eggs rather than by widening the template for acceptable
queen-laid eggs. These novel results show that egg-discrimination behaviour in F. fusca is flexible and that workers respond to the diversity of eggs experienced in their colony. 相似文献
9.
Insect societies are sometimes exploited by workers who reproduce selfishly rather than help to rear the queens offspring. This causes a conflict-of-interest with the mother queen and, frequently, with the non-reproductive workers as well. One mechanism that can reduce conflict is policing, whereby either the queen or other workers aggress egg-laying workers or destroy worker-laid eggs. Here we present the first direct observations of queen and worker policing in natural, unmanipulated colonies of a social insect, the tree wasp Dolichovespula sylvestris. Worker reproduction was common, with workers producing 50% of all male eggs. However, most worker-laid eggs, 91%, were policed within 1 day, whereas most queen-laid eggs, 96%, remained unharmed. The workers were responsible for 51% of all policing events and the queen for 49%. The workers and mother queen also commonly aggressed ovipositing workers, and successfully prevented them from depositing eggs in 14% and 6% of all attempted ovipositions. Hence, both queen policing and worker policing occur and policing acts via two distinct mechanisms: selective destruction of worker-laid eggs and aggression of ovipositing workers. At a general level, our study shows that both centralized and decentralized control can act together to suppress conflict within social groups.Communicated by R. Page 相似文献
10.
P. Kirk Visscher 《Behavioral ecology and sociobiology》1996,38(4):237-244
Using electrophoretic markers, eggs laid by workers were identified in honey bee (Apis mellifera) colonies with a queen. Based on extrapolation, these represented about 7% of the unfertilized (male) eggs laid in the colonies.
A very small proportion of workers (of the order of 0.01%) lay these eggs. Worker-laid eggs are rapidly removed, so that very
few sons of workers are reared. Thus the reproductive cooperation in bee colonies is maintained by ongoing antagonistic interactions
among the members of the colony, with worker laying and egg removal policing by other workers being relatively common.
Received: 24 November 1995/Accepted after revision: 25 May 1996 相似文献
11.
Shelley E. R. Hoover Mark L. Winston Benjamin P. Oldroyd 《Behavioral ecology and sociobiology》2005,59(2):278-284
In most social insect colonies, workers do not attempt to lay eggs in the presence of a queen. However, in the honey bee (Apis mellifera), a rare phenotype occurs in which workers activate their ovaries and lay large numbers of male eggs despite the presence of a fecund queen. We examined the proximate mechanisms by which this ‘anarchistic’ behaviour is expressed. We tested the effects of brood and queen pheromones on retinue attraction and worker ovary activation using caged worker bees. We found no difference between the anarchistic and wild type queen pheromones in the retinue response elicited in either wild type or anarchistic workers. Further, we found that anarchistic queens produce a pheromone blend that is as effective at inhibiting ovary activation as the wild type queen pheromone. However, anarchistic workers are less inhibited by queen pheromones than their wild type counterparts, in a dose-dependent manner. These results show that the anarchistic phenomenon is not due to changes in the production of queen pheromones, but rather is due in part to a shift in the worker response to these queen-produced signals. In addition, we demonstrate the dose-dependent nature of the effect of queen pheromones on honey bee worker ovary activation. 相似文献
12.
Luke A. Halling Benjamin P. Oldroyd Wandee Wattanachaiyingcharoen Andrew B. Barron Piyamas Nanork Siriwat Wongsiri 《Behavioral ecology and sociobiology》2001,49(6):509-513
Apis florea is a single-combed, open-nesting, dwarf honeybee indigenous to Asia. In common with other species of this genus, A. florea is highly polyandrous, and is therefore predicted to curtail worker reproduction by mutual policing mechanisms that keep worker reproduction at an extremely low level. Policing mechanisms could involve destruction of workers' eggs or offspring, or aggression toward those workers that are reproductively active. We show that in A. florea, worker-laid eggs are eliminated approximately twice as fast as queen-laid eggs, indicating that A. florea uses oophagy of worker-laid eggs as a mechanism of worker policing. Genetic analysis of four colonies indicated that all males produced were sons of queens, not workers. Dissections of 800 workers, from four colonies, did not reveal any significant levels of ovary activation. These results suggest that worker policing is an effective component of the mechanisms that maintain worker sterility in this species. Furthermore, they suggest that worker policing via oophagy of worker-laid eggs is pleisiomorphic for the genus. 相似文献
13.
Madeleine Beekman Stephen J. Martin Falko Drijfhout Benjamin P. Oldroyd 《Behavioral ecology and sociobiology》2007,61(12):1847-1853
Anarchy, where honeybee workers produce males in the presence of a queen, is extremely rare in natural honeybee populations,
suggesting that there are colony-level costs associated with being anarchistic. Yet, no significant costs have yet been identified.
A possible reason for this may be that researchers have only focused on the behaviour of anarchistic workers, which have been
shown to perform worker-tasks as do wild-type workers. Possibly, therefore, costs associated with anarchy should be sought
in anarchistic queens and not workers. A potential cost could be a lower survival rate of eggs laid by anarchistic queens
perhaps because their egg-marking signal is not as clear as that of a wild-type queen. In this paper, we determined the removal
rate of eggs laid by anarchistic queens in standard worker-policing bioassays. Our results show that eggs laid by anarchistic
queens are removed at a higher rate than eggs laid by wild-type queens. This does not seem to be due to differences in hydrocarbons
found on the surface of eggs, as both egg types showed the same alkanes and alkenes in similar proportions. We postulate that
higher removal rates of queen-laid eggs due to recognition errors may be one reason that anarchy is rare in natural honeybee
populations. 相似文献
14.
The Red Dwarf honeybee (Apis florea) is one of two basal species in the genus Apis. A. florea differs from the well-studied Western Hive bee (Apis mellifera) in that it nests in the open rather than in cavities. This fundamental difference in nesting biology is likely to have implications
for nest-site selection, the process by which a reproductive swarm selects a new site to live in. In A. mellifera, workers show a series of characteristic behaviors that allow the swarm to select the best nest site possible. Here, we describe
the behavior of individual A. florea workers during the process of nest-site selection and show that it differs from that seen in A. mellifera. We analyzed a total of 1,459 waggle dances performed by 197 scouts in five separate swarms. Our results suggest that two
fundamental aspects of the behavior of A. mellifera scouts—the process of dance decay and the process of repeated nest site evaluation—do not occur in A. florea. We also found that the piping signal used by A. mellifera scouts to signal that a quorum has been reached at the chosen site, is performed by both dancing and non-dancing bees in
A. florea. Thus, the piping signal appears to serve a different purpose in A. florea. Our results illustrate how differences in nesting biology affect the behavior of individual bees during the nest-site selection
process. 相似文献
15.
Margaret J. Couvillon Francis L.W. Ratnieks 《Behavioral ecology and sociobiology》2008,62(7):1099-1105
In group-level recognition, discriminators use sensory information to distinguish group members and non-members. For example,
entrance guards in eusocial insect colonies discriminate nestmates from intruders by comparing their odour with a template
of the colony odour. Despite being a species-rich group of eusocial bees closely related to the honey bees, stingless bee
nestmate recognition is a relatively little-studied area. We studied Frieseomelitta varia, a common Brazilian species of stingless bee known as marmelada. By measuring the rejection rates of nestmate and non-nestmate
worker bees by guards, we were able to show that guards became significantly less accepting (from 91 to 46%) of nestmates
that had acquired odour cues from non-nestmate workers; however, guards did not become significantly more accepting (from
31 to 42%) of non-nestmates that had acquired equivalent amounts of odour cues from the guard’s nestmates. These data strongly
suggest that guards use an “undesirable–absent” system in recognition, whereby incoming conspecific workers are only accepted
if undesirable cues are absent, despite the presence of desirable cues. We suggest that an undesirable–absent system is adaptive
because robbing by conspecifics may be an important selective factor in F. varia, which would lead to selection for a non-permissive acceptance strategy by guards. 相似文献
16.
Piyamas Nanork Siriwat Wongsiri Benjamin P. Oldroyd 《Behavioral ecology and sociobiology》2006,61(1):91-97
Honeybee (Apis) workers cannot mate, but retain functional ovaries. When colonies have lost their queen, many young workers begin to activate their ovaries and lay eggs. Some of these eggs are reared, but most are not and are presumably eaten by other workers (worker policing). Here we explore some of the factors affecting the reproductive success of queenless workers of the red dwarf honeybee Apis florea. Over a 2-year period we collected 40 wild colonies and removed their queens. Only two colonies remained at their translocated site long enough to rear males to pupation while all the others absconded. Absconding usually occurred after worker policing had ceased, as evidenced by the appearance of larvae. Dissections of workers from eight colonies showed that in A. florea, 6% of workers have activated ovaries after 4 days of queenlessness, and that 33% of workers have activated ovaries after 3 weeks. Worker-laid eggs may appear in nests within 4 days and larvae soon after, but this is highly variable. As with Apis mellifera, we found evidence of unequal reproductive success among queenless workers of A. florea. In the two colonies that reared males to pupation and which we studied with microsatellites, some subfamilies had much higher proportions of workers with activated ovaries than others. The significance of absconding and internest reproductive parasitism to the alternative reproductive strategies of queenless A. florea workers is discussed. 相似文献
17.
Michael H. Allsopp Madeleine Beekman Rosalyn S. Gloag Benjamin P. Oldroyd 《Behavioral ecology and sociobiology》2010,64(4):567-574
Unlike workers of all other honey bee (Apis mellifera) subspecies, workers of the Cape honey bee of South Africa (A. mellifera capensis) reproduce thelytokously and are thus able to produce female offspring that are pseudoclones of themselves. This ability
allows workers to compete with their queen over the maternity of daughter queens and, in one extreme case, has led to a clonal
lineage of workers becoming a social parasite in commercially managed populations of A. mellifera scutellata. Previous work (Jordan et al., Proc R Soc Lond B Biol Sci 275:345, 2008) showed that, in A. mellifera capensis, 59% of queen cells produced during swarming events contained the offspring of workers and that, of these, 65% were the offspring
of non-natal workers. Here, we confirm that a substantial proportion (38.5%) of offspring queens is worker-laid. We additionally
show that: (1) Although queens produce most diploid female offspring sexually, we found some homozygous or hemizygous queen
offspring, suggesting that queens also reproduce by thelytoky. These parthenogenetic individuals are probably nonviable beyond
the larval stage. (2) Worker-laid offspring queens are viable and become the resident queen at the same frequency as do sexually
produced queen-laid offspring queens. (3) In this study, all but one of the worker-derived queens were laid by natal workers
rather than workers from another nest. This suggests that the very high rates of social parasitism observed in our previous
study were enhanced by beekeeping manipulations, which increased movement of parasites between colonies. 相似文献
18.
Joël Meunier Luma Delaplace Michel Chapuisat 《Behavioral ecology and sociobiology》2010,64(10):1655-1663
The ability to discriminate against competitors shapes cooperation and conflicts in all forms of social life. In insect societies,
workers may detect and destroy eggs laid by other workers or by foreign queens, which can contribute to regulate reproductive
conflicts among workers and queens. Variation in colony kin structure affects the magnitude of these conflicts and the diversity
of cues used for discrimination, but the impact of the number of queens per colony on the ability of workers to discriminate
between eggs of diverse origin has so far not been investigated. Here, we examined whether workers from the socially polymorphic
ant Formica selysi distinguished eggs laid by nestmate workers from eggs laid by nestmate queens, as well as eggs laid by foreign queens from
eggs laid by nestmate queens. Workers from single- and multiple-queen colonies discriminated worker-laid from queen-laid eggs,
and eliminated the former. This suggests that workers collectively police each other in order to limit the colony-level costs
of worker reproduction and not because of relatedness differences towards queens’ and workers’ sons. Workers from single-queen
colonies discriminated eggs laid by foreign queens of the same social structure from eggs laid by nestmate queens. In contrast,
workers from multiple-queen colonies did not make this distinction, possibly because cues on workers or eggs are more diverse.
Overall, these data indicate that the ability of F. selysi workers to discriminate eggs is sufficient to restrain worker reproduction but does not permit discrimination between matrilines
in multiple-queen colonies. 相似文献
19.
Willem J. Boot Minus van Baalen Maurice W. Sabelis 《Behavioral ecology and sociobiology》1995,36(4):283-289
Varroa jacobsoni reproduces both in drone and worker brood cells of honey bees, but in drone cells reproductive success is higher than in worker cells. A simple model using clonal population growth as a fitness measure has been developed to study the circumstances under which specialization on drone brood would be a better strategy than reproduction in both types of cell. For European Apis mellifera, the model suggests that if mites have to wait less than 7 days on average before they can invade a drone cell, specialization on drone brood would be a better strategy. This is close to the estimated waiting time of 6 days. Hence, small differences in reproductive success in drone and worker cells and in the rate of mortality may determine whether specialization on drone brood will be promoted or not. In European A. mellifera colonies, Varroa mites invade both drone and worker cells, but specialization on drone brood cells seems to occur to some extent because drone cells are more frequently invaded than worker cells. In the parasite-host association of V. jacobsoni with African or Africanized A. mellifera or with A. cerana, the mites also invade both drone and worker cells, but the mites specialize on drone brood for reproduction since a large percentage of the mites in worker brood do not reproduce. Only in the parasite-host association of Euvarroa sinhai, a mite closely resembling V. jacobsoni, and A. forea is specialization complete, because these mites only invade drone brood. 相似文献
20.
With very rare exceptions, queenright worker honeybees (Apis mellifera L.) forego personal reproduction and suppress reproduction by other workers, preferring to rear the queens sons. This is in stark contrast to colonies that have lost their queen and have failed to rear a replacement. Under these conditions workers activate their ovaries and lay many eggs that develop parthenogenetically into a final brood of males (drones) before the colony perishes. Interestingly, not all workers contribute equally to this final generation of drones in queenless colonies. Some subfamilies (workers that share the same father) contribute a disproportionately greater number of offspring than other subfamilies. Here we explore some of the mechanisms behind this reproductive competition among subfamilies. We determined the relative contribution of different subfamilies present in colonies to laying workers, eggs, larvae and pupae by genotyping samples of all life stages using a total of eight microsatellite loci. Our colonies were headed by free-mated queens and comprised 8–17 subfamilies and therefore differed significantly from colonies used in an earlier study investigating the same phenomena where colonies comprised an artificially low number of subfamilies. We show that, first, subfamilies vary in the speed with which they activate their ovaries after queen-loss and, second, that the survival of eggs to the larval stage is unequal among subfamilies suggesting that some subfamilies lay eggs that are more acceptable than others. However, there is no statistically significant difference among subfamilies in the survival of larvae to pupae, indicating that ovary activation and egg survival are the critical components to reproductive competition among subfamilies of queenless honeybee workers.Communicated by R. Page 相似文献