Phenotype interactions in group behavior of honey bee workers (Apis mellifera L.)

Summary The alarm reaction of groups of honey bee workers was quantified using a metabolic bioassay. The genetic structure of these groups was varied in order to estimate the effects of worker interactions. Though the group phenotype was mainly determined by additive interactions, nonlinear effects were also found. Mixed worker groups, combined from colonies with similar reactivity in the bioassay, showed a stronger response than pure groups. This phenomenon, analogous to the overdominance model for individuals in classical genetics, has implications for mechanisms of natural and artificial selection in social populations and for the evolution of polyandry in social Hymenoptera.     

Social parasitism by honeybee workers (Apis mellifera capensis Esch.): evidence for pheromonal resistance to host queen’s signals

Social parasites exploit their host’s communication system to usurp resources and reproduce. In the honeybee, Apis mellifera, worker reproduction is regulated by pheromones produced by the queen and the brood. Workers usually reproduce when the queen is removed and young brood is absent. However, Cape honeybee workers, Apis mellifera capensis, are facultative intraspecific social parasites and can take over reproduction from the host queen. Investigating the manner in which parasitic workers compete with host queens pheromonally can help us to understand how such parasitism can evolve and how reproductive division of labour is regulated. In A. m. capensis, worker reproduction is associated with the production of queen-like pheromones. Using pheromonal contest experiments, we show that Apis mellifera scutellata queens do not prevent the production of queen-like mandibular gland compounds by the parasites. Given the importance of these pheromones in acquiring reproductive status, our data suggest that the single invasive lineage of parasitic workers occurring in the range of A. m. scutellata was selected for its superior ability to produce these signals despite the presence of a queen. Such resistance was indeed less frequent amongst other potentially parasitic lineages. Resistance to reproductive regulation by host queens is probably the key factor that facilitates the evolution of social parasitism by A. m. capensis workers. It constitutes a mechanism that allows workers to evade reproductive division of labour and to follow an alternative reproductive option by acquiring direct fitness in foreign colonies instead of inclusive fitness in their natal nests.     

Ovariole number—a predictor of differential reproductive success among worker subfamilies in queenless honeybee (Apis mellifera L.) colonies

A honeybee queen normally mates with 10–20 drones, and reproductive conflicts may arise among a colony’s different worker patrilines, especially after a colony has lost its single queen and the workers commence egg laying. In this study, we employed microsatellite markers to study aspects of worker reproductive competition in two queenless Africanized honeybee colonies. First, we determined whether there was a bias among worker patrilines in their maternity of drones and, second, we asked whether this bias could be attributed to differences in the degree of ovary activation of workers. Third, we relate these behavioral and physiological factors to ontogenetic differences between workers with respect to ovariole number. Workers from each of three (colony A) and one (colony B) patrilineal genotypes represented less than 6% of the worker population, yet each produced at least 13% of the drones in a colony, and collectively they produced 73% of the drones. Workers representing these genotypes also had more developed follicles and a greater number of ovarioles per ovary. Across all workers, ovariole development and number were closely correlated. This suggests a strong effect of worker genotype on the development of the ovary already in the postembryonic stages and sets a precedent to adult fertility, so that “workers are not born equal”. We hypothesize a frequency-dependent or “rare patriline” advantage to queenless workers over the parentage of males and discuss the maintenance of genetic variance in the reproductive capacity of workers.Electronic supplementary material Supplementary material is available for this article at and is accessible for authorized users.     

Differential reproductive success among subfamilies in queenless honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.) colonies

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     

The influence of pollen quality on foraging behavior in honeybees (Apis mellifera L.)

Individual and colony-level foraging behaviors were evaluated in response to changes in the quantity or nutritional quality of pollen stored within honeybee (Apis mellifera L.) colonies. Colonies were housed in vertical, three-frame observation hives situated inside a building, with entrances leading to the exterior. Before receiving treatments, all colonies were deprived of pollen for 5 days and pollen foragers were marked. In one treatment group, colony pollen reserves were quantitatively manipulated to a low or high level, either by starving colonies of pollen or by providing them with a fully provisioned frame of pollen composed of mixed species. In another treatment group, pollen reserves were qualitatively manipulated by removing pollen stores from colonies and replacing them with low- or high-protein pollen supplements. After applying treatments, foraging rates were measured four times per day and pollen pellets were collected from experienced and inexperienced foragers to determine their weight, species composition, and protein content. Honeybee colonies responded to decreases in the quantity or quality of pollen reserves by increasing the proportion of pollen foragers in their foraging populations, without increasing the overall foraging rate. Manipulation of pollen stores had no effect on the breadth of floral species collected by colonies, or their preferences for the size or protein content of pollen grains. In addition, treatments had no effect on the weight of pollen loads collected by individual foragers or the number of floral species collected per foraging trip. However, significant changes in foraging behavior were detected in relation to the experience level of foragers. Irrespective of treatment group, inexperienced foragers exerted greater effort by collecting heavier pollen loads and also sampled their floral environment more extensively than experienced foragers. Overall, our results indicate that honeybees respond to deficiencies in the quantity or quality of their pollen reserves by increasing the gross amount of pollen returned to the colony, rather than by specializing in collecting pollen with a greater protein content. Individual pollen foragers appear to be insensitive to the quality of pollen they collect, indicating that colony-level feedback is necessary to regulate the flow of protein to and within the colony. Colonies may respond to changes in the quality of their pollen stores by adjusting the numbers of inexperienced to experienced foragers within their foraging populations.     

The effects of colony-level selection on the social organization of honey bee (Apis mellifera L.) colonies: colony-level components of pollen hoarding

Two-way selection for quantities of stored pollen resulted in the production of high and low pollen hoarding strains of honey bees (Apis mellifera L.). Strains differed in areas of stored pollen after a single generation of selection and, by the third generation, the high strain colonies stored an average 6 times more pollen than low strain colonies. Colony-level organizational components that potentially affect pollen stores were identified that varied genetically within and between these strains. Changes occurred in several of these components, in addition to changes in the selected trait. High strain colonies had a significantly higher proportion of foragers returning with loads of pollen, however, high and low strain colonies had equal total numbers of foragers Colony rates of intake of pollen and nectar were not independent. Selection resulted in an increase in the number of pollen collectors and a decrease in the number of nectar collectors in high strain colonies, while the reciprocal relationship occurred in the low strain. High and low strain colonies also demonstrated different diurnal foraging patterns as measured by the changing proportions of returning pollen foragers. High strain colonies of generation 3 contained significantly less brood than did low strain colonies, a consequence of a constraint on colony growth resulting from a fixed nest volume and large quantities of stored pollen. These components represent selectable colony-level traits on which natural selection can act and shape the social organization of honey bee coloniesCommunicated by R.F.A. Moritz     

Social parasitism of queens and workers in the Cape honeybee (<Emphasis Type="Italic">Apis mellifera capensis</Emphasis>)

Workers of a queenless honeybee colony can requeen the colony by raising a new queen from a young worker brood laid by the old queen. If this process fails, the colony becomes hopelessly queenless and workers activate their ovaries to lay eggs themselves. Laying Cape honeybee workers (Apis mellifera capensis) produce female offspring as an additional pathway for requeening. We tested the frequency of successful requeening in ten hopelessly queenless colonies. DNA genotyping revealed that only 8% of all queens reared in hopelessly queenless colonies were the offspring of native laying worker offspring. The vast majority of queens resulted from parasitic takeovers by foreign queens (27%) and invading parasitic workers (19%). This shows that hopelessly queenless colonies typically die due to parasitic takeovers and that the parasitic laying workers are an important life history strategy more frequently used than in providing a native queen to rescue the colony. Parasitism by foreign queens, which might enter colonies alone or accompanied by only a small worker force is much more frequent than previously considered and constitutes an additional life history strategy in Cape honeybees.     

Self-organization of circadian rhythms in groups of honeybees (Apis mellifera L.)

Workers in social groups of honeybees (Apis mellifera L.) synchronize their individual free-running circadian rhythms to an overall group rhythm. By monitoring the activity of bees by recording the oxygen consumption and intragroup temperature, it is shown that the rhythm coordination is in part achieved by temperature fluctuations as an intragroup Zeitgeber. Trophallaxis was shown to have only a minor (if any) effect on circadian rhythm synchronization. A model incorporating a feed back loop between temperature and activity can plausibly explain the observed synchronization of individual rhythms in social groups as a self-organization phenomenon. Correspondence to: R.F.A. Moritz     

Optimal timing of comb construction by honeybee (Apis mellifera) colonies: a dynamic programming model and experimental tests

Honeybee colonies, like organisms, should exhibit optimal design in their temporal pattern of resource allocation to somatic structures. A vital colony structure is the comb which stores honey for overwinter survival. However, the timing of comb construction poses a dilemma to a colony attempting to maximize its honey reserves. On the one hand, plenty of empty comb is needed for efficient exploitation of temporally unpredictable flower blooms. On the other hand, because comb is made from energetically expensive wax, its construction too early or in excessive amounts will reduce the amount of honey available for winter thermoregulation and brood-rearing. A dynamic optimization model concludes that colonies should add new comb only when they have filled their old comb with food and brood above a threshold level. The threshold increases with time until, at the end of the season, building is never an optimal behavior. The temporal pattern of construction predicted by the model – pulses of building coincident with periods of nectar intake and comb fullness – matches that seen in an actual colony observed over the course of an entire foraging season. When nectar sources are rich but temporally clumped, the model also predicts that bees should be sensitive to nectar intake, employing much higher thresholds on days when nectar is not available than on days when it is. Even under poorer and more dispersed nectar regimes, little fitness cost is paid by colonies replacing the optimal strategy with a simpler rule of thumb calling for new construction only when two conditions are met: (1) a fullness threshold has been exceeded, and (2) nectar is currently being collected. Experiments demonstrate that colonies do in fact use such a rule of thumb to control the onset of construction. However, once they have begun building, the bees continue as long as nectar collection persists, regardless of changes in comb fullness. Thus the onset and duration of comb-building bouts appear to be under partially independent control. Received: 30 October 1998 / Received in revised form: 14 December 1998 / Accepted: 16 January 1999     

Sperm limitation and the evolution of extreme polyandry in honeybees (Apis mellifera L.)

Honeybee queens (Apis mellifera) show extreme levels of polyandry, but the evolutionary mechanisms underlying this behaviour are still unclear. The sperm-limitation hypothesis, which assumes that high levels of polyandry are essential to get a lifetime sperm supply for large and long-lived colonies, has been widely disregarded for honeybees because the semen of a single male is, in principle, sufficient to fill the spermatheca of a queen. However, the inefficient post-mating sperm transfer from the queens lateral oviducts into the spermatheca requires multiple matings to ensure an adequate spermatheca filling. Males of the African honeybee subspecies A. m. capensis have fewer sperm than males of the European subspecies A. m. carnica. Thus, given that sperm limitation is a cause for the evolution of multiple mating in A. mellifera, we would expect A. m. capensis queens to have higher mating frequencies than A. m. carnica. Here we show that A. m. capensis queens indeed exhibit significantly higher mating frequencies than queens of A. m. carnica, both in their native ranges and in an experiment on a North Sea island under the same environmental conditions. We conclude that honeybee queens try to achieve a minimum number of matings on their mating flights to ensure a sufficient lifetime sperm supply. It thus seems premature to reject the sperm-limitation hypothesis as a concept explaining the evolution of extreme polyandry in honeybees.Communicated by R.E. Page     

Colony performance in honeybees (Apis mellifera capensis Esch.) depends on the proportion of subordinate and dominant workers

Summary Individual worker dominance correlated with trophallactic behavior, which affects several social behaviors related to colony fitness, shows a high genetic variance in worker bees. In a bioassay we tested trophallactic behavior of workers and selected dominant (receiving) and subordinate worker bees (offering) of Apis mellifera capensis to establish genetic lines of both kinds. Queenright test colonies were experimentally composed of 100% subordinate workers, 100% dominant workers, 50% dominant plus 50% subordinate workers, and 100% hybrid workers from the two genetic lines. The chosen test parameters were brood-rearing, comb building and hoarding behavior. In all cases, the colonies of pure subordinate bees showed the best colony performance, whereas the colonies composed of only dominant bees were nearly unproductive. The mixed colonies (50% dominant + 50% subordinate) ranked in the middle and did not differ significantly from the hybrid colonies. The results indicate that colony performance under queenright conditions depends on the proportion of subordinate workers. This result supports a selection model based on the combination of individual selection and on group selection at the colony level, which explains the high genetic variance of individual worker reproduction.     

Behavioral and life-history components of division of labor in honey bees (Apis mellifera L.)

Variability exists among worker honey bees for components of division of labor. These components are of two types, those that affect foraging behavior and those that affect life-history characteristics of workers. Variable foraging behavior components are: the probability that foraging workers collect (1) pollen only; (2) nectar only; and (3) pollen and nectar on the same trip. Life history components are: (1) the age the workers initiate foraging behavior; (2) the length of the foraging life of a worker; and (3) worker length of life. We show how these components may interact to change the social organization of honey bee colonies and the lifetime foraging productivity of individual workers. Selection acting on foraging behavior components may result in changes in the proportion of workers collecting pollen and nectar. Selection acting on life-history components may affect the size of the foraging population and the distribution of workers between within nest and foraging activities. We suggest that these components define possible sociogenic pathways through which colony-level natural selection can change social organization. These pathways may be analogous to developmental pathways in the morphogenesis of individual organisms because small changes in behavioral or life history components of individual workers may lead to major changes in the organizational structure of colonies. Correspondence to: R.E. Page, Jr.     

Effects of worker genotypic diversity on honey bee colony development and behavior (Apis mellifera L.)

There have been numerous reports of genetic influences on division of labor in honey bee colonies, but the effects of worker genotypic diversity on colony behavior are unclear. We analyzed the effects of worker genotypic diversity on the phenotypes of honey bee colonies during a critical phase of colony development, the nest initiation phase. Five groups of colonies were studied (n = 5–11 per group); four groups had relatively low genotypic diversity compared to the fifth group. Colonies were derived from queens that were instrumentally inseminated with the semen of four different drones according to one of the following mating schemes: group A, 4 A-source drones; group B, 4 B-source drones; group C, 4 C-source drones; group D, 4 D-source drones; and group E, 1 drone of each of the A-D drone sources. There were significant differences between colonies in groups A-D for 8 out of 19 colony traits. Because the queens in all of these colonies were super sisters, the observed differences between groups were primarily a consequence of differences in worker genotypes. There were very few differences (2 out of 19 traits) between colonies with high worker genotypic diversity (group E) and those with low diversity (groups A-D combined). This is because colonies with greater diversity tended to have phenotypes that were average relative to colonies with low genotypic diversity. We hypothesize that the averaging effect of genotypic variability on colony phenotypes may have selective advantages, making colonies less likely to fail because of inappropriate colony responses to changing environmental conditions.     

Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

三唑磷降解对水稻叶绿素含量的影响

采用田间试验方法,系统研究了内2优6号、秀水09两个水稻品种生殖生长前期经过喷施不同浓度三唑磷(2 250 mL hm-2、4 500 mL hm-2)后水稻叶绿素含量动态变化及其相关性.结果表明,喷施三唑磷对水稻叶绿素含量产生了影响,其效应随施药浓度增加而增大.三唑磷对叶绿素作用的时效在品种间存在差异,内2优6号和秀水09分别为<20 d和>20 d,具体表现为施药后d 1,所有水稻品种的喷施处理与对照无显著性差异,施药后d 3～14,与对照相比内2优6号和秀水09分别表现为显著降低和增加,施药后d 21,则分别表现为不显著和显著增加.三唑磷降解率与叶绿素含量变化率之间总体存在相关性,但品种间存在显著差异,内2优6号和秀水09分别呈现负相关和正相关趋势.     

沙芥幼苗叶片解剖结构和光合作用对干旱胁迫的响应

为了了解干旱胁迫对植物光合作用和叶片结构的影响,明确植物光合作用和结构对干旱胁迫的响应,本研究以沙生植物、中国特有种---沙芥为材料,采用Li-6400光合仪和常规石蜡切片法,研究了干旱胁迫下沙芥光合参数、叶片解剖结构的变化,结合前期对沙芥叶片内活性氧物质含量变化的研究,试图阐述沙芥叶片应对干旱胁迫的机制,揭示沙生植物—沙芥在干旱环境中的生存策略,为植物耐旱性提供理论依据。结果表明：随着土壤含水量下降,净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、叶片厚度、上表皮厚度、下表皮厚度呈下降趋势;海绵组织厚度呈先下降后上升的趋势;胞间二氧化碳浓度(Ci)呈上升的趋势;气孔限制值(Ls)、水分利用率(WUE)、栅栏组织和栅栏组织/海绵组织和叶片组织结构紧密度(CTR)呈先先上升后下降的趋势;且在土壤含水量大于39% WHC时沙芥光合和结构受影响较小,在土壤含水量低于39% WHC时,沙芥光合作用受到抑制,而叶片紧密度变大,能够较好的抵御干旱环境。结果表明干旱胁迫下沙芥幼苗叶片光合作用的下降是非气孔因素造成的,且其叶片结构的变化会影响光合作用,光合作用通过产生活性氧对结构也有影响。说明沙芥能够很好的适应中度干旱以上的环境,在重度干旱环境中沙芥通过调节叶片结构以适应环境,这可能是沙芥能够在沙漠地区生存繁殖的原因之一。     

镉胁迫对水车前叶片抗氧化酶系统和亚显微结构的影响

报道了模拟镉污染对水车前叶片抗氧化酶系统的影响及对叶细胞超微结构 ,主要是叶绿体、线粒体和细胞核的损伤情况。随胁迫程度的增大 ,超氧化物歧化酶活性和可溶性蛋白含量下降明显 ,丙二醛含量则上升 ,过氧化物酶和过氧化氢酶活性在 0 1mg/L浓度处达到最高后呈下降趋势。电镜观察结果表明 :毒害初期 ,叶绿体和线粒体膨胀 ,类囊体明显空泡化 ;高浓度镉处理使破坏程度加重 ,叶绿体被膜消失和叶绿体解体 ;线粒体间质溶解 ,濒临解体 ;细胞核染色质凝集并散入细胞质中。由于镉胁迫破坏了植物的抗氧化酶系统 ,并对超微结构造成不可逆损伤 ,最终使细胞死亡     

干旱区哈密瓜生长和产量与气候要素的关系

农业是气候变化最敏感的领域之一,气候变化将对农作物产量产生巨大影响.探讨气候变化对干旱区哈密瓜(Cucumis melo L.)生长和产量的影响,明确关键影响气象因子,可为合理有效利用气候资源和防灾减灾提供科学依据.选取1981—2019年哈密瓜生长年气候因子、生长季内气象要素和1999—2019年发育期及产量资料,分...     

Comparisons in subcellular and biochemical behaviors of cadmium between Iow-Cd and high-Cd accumulation cultivars of pakchoi （Brassica chinensis L.）

Subcellular distributions and chemical forms of cadmium （Cd） in the leaves, stems and roots were investigated in low-Cd accumulation cultivars and high-Cd accumulation cultivars ofpakchoi （Brassica chinensis L.）. Root cell wall played a key role in limiting soil Cd from entering the protoplast, especially in the low-Cd cultivars. The high-Cd cultivars had significantly higher leaf and stem Cd concentrations than the low-Cd cultivars in cell wall fraction, chloroplast/trophoplast fraction, organelle fraction and soluble fraction. In low-Cd cultivars, which were more sensitive and thus had greater physiological needs of Cd detoxification than high-Cd cultivars, leaf vacuole sequestrated higher proportions of Cd. Cd in the form of pectate/protein complexes （extracted by 1 tool. L~ NaC1） played a decisive role in Cd translocation from root to shoot, which might be one of the mechanisms that led to the differences in shoot Cd accumulation between the two types of cultivars. Furthermore, the formation of Cd- phosphate complexes （extracted by 2% HAc） was also involved in Cd detoxification within the roots of pakchoi under high Cd stress, suggesting that the mechanisms of Cd detoxification might be different between low- and high-Cd cultivars.