明德译介 | 蜂群及其组织

原文:The Colony and Its Organization
编译:沙皮狗

蜜蜂是群居性昆虫,这意味着它们共同生活在一个组织良好的大型家庭群体中。群居昆虫是高度进化的昆虫,它们从事多种独居昆虫无法完成的复杂任务。沟通、复杂的巢穴建造、环境控制、防御和分工是蜜蜂已经发展到在社会群体中成功存在的其中一些行为。这些的行为使社会性昆虫,特别是蜜蜂,成为地球上最让人著迷的生物之一。

蜜蜂群通常由三种成年蜜蜂组成:工蜂、雄蜂和蜂后。数千只工蜂在筑巢、收集食物和育雏方面合作。每个成员都有与其成年年龄相关的明确任务要执行。但生存和繁衍需要整个蜂群的共同努力。没有蜂群的支持,个体蜜蜂(工蜂、雄蜂和蜂后)无法生存。

除了成千上万的成年工蜂外,一个蜂群通常在春末和夏季有一个蜂后和数百架雄蜂。蜂群的社会结构由蜂后和工人的存在维持,并依赖于有效的沟通系统。化学信息素在成员之间的分布和交流的“舞蹈”负责控制蜂群生存所必需的活动。工蜂的劳动活动主要取决于蜜蜂的年龄,但会随着蜂群的需要而变化。繁殖和蜂群的力量取决于蜂后、食物的数量和劳动力的规模。随着蜂群的规模增加到最多约 60,000 名工蜂,蜂群的效率也随之提高。

蜂后

每个蜂群只有一个蜂后---除了在蜂群准备分群或取代老弱蜂后的时期。因为她是唯一性发育成熟的女性,所以她的主要功能是繁殖。她生产受精卵和未受精卵。蜂后在春季和初夏产卵最多。在生产高峰期,蜂后每天可产下多达 1,500 个卵。它们在 10 月初逐渐停止产卵,直到明年春天(1 月)初产卵或很少产卵。一蜂后每年可产卵多达 250,000 个,一生中可能超过 100 万个。

蜂后很容易与蜂群的其他成员区分开来。她的身体通常比雄蜂或工人的身体长得多,尤其是在产卵期间,她的腹部大大拉长。她的翅膀只覆盖了腹部的三分之二左右,而工蜂和雄蜂的翅膀在折叠时几乎达到了腹部的尖端。蜂后的胸部比工蜂的略大,她既没有花粉篮也没有功能性的蜡腺。她的毒刺是弯曲的,比工人的长,但它的倒刺更少更短。蜂后可以活几年——有时长达 5 年,但平均生产寿命为 2 到 3 年。

蜂后的第二个主要功能是产生信息素,作为一种社会“粘合剂”,统一并帮助赋予蜂群个体身份。一种主要的信息素——称为蜂后物质——是由她的下颌腺产生的,但其他的也很重要。蜂群的品质很大程度上取决于蜂后的产卵和化学生产能力。她的基因构成——以及她与之交配的雄蜂的基因构成——对群体的质量、大小和行为产生重大影响。

从蜂后房出来大约一周后,蜂后离开蜂巢与飞行中的几只雄蜂交配。因为她必须从她的蜂群飞离一段距离才能交配(大自然避免近亲繁殖的方法),所以她首先绕着蜂巢绕圈定位自己的位置。她独自离开蜂巢,大约 13 分钟后离开。蜂后通常在下午与 7 到 15 只雄蜂在 20 英尺以上的高度交配。雄蜂能够通过她的化学气味(信息素)找到并识别蜂后。如果恶劣的天气使蜂后的交配出行延误超过 20 天,她就会失去交配能力,只能产下未受精的卵,从而产生雄峰。

交配后,蜂后返回蜂巢并在大约 48 小时内开始产卵。每次她都会从受精囊中释放出几个精子,这会注定她产下一个的是成为工人或蜂后的卵。如果她的卵产在一个较大的育蜂室(雄蜂用)时,她就不会释放精子。蜂后经常被蜂群的工蜂照料并喂食蜂皇浆。蜂后产卵的数量取决于她收到的食物量以及准备了多少用蜂蜡做的育蜂室和照顾将在 3 天内从卵中孵化出来的幼虫的劳动力规模。当蜂后分泌的蜂后物质不再充足时,工人们准备替换(取代)她。老蜂后和她的新女儿可能在被取代后都在蜂巢中同时存在一段时间。

新的(处女)皇后由受精卵或不超过 3 天大的工蜂幼虫发育而成。新蜂后在三种不同的情况下产生:紧急情况、取代老蜂后或蜂群分家。当一只老蜂后被意外杀死、丢失或移走时,蜜蜂会选择年轻的工蜂幼虫来生产应急蜂后。这些蜂后在经过修改以垂直悬挂在梳子表面上的工蜂育蜂室中长大。当一个蜂后开始衰老(减少蜂后物质的产量)时,蜂群准备培养一个新的蜂后。因替代而产生的蜂后通常比紧急蜂后好,因为它们在发育过程中会接受更多的食物(蜂后浆)。与紧急蜂后育蜂室一样,替代蜂后育蜂室通常在梳状表面上凸起。相比之下,为蜂群准备而产生的蜂后育蜂室位于框架的底部边缘或育雏区域内蜂蜡梳的间隙中。

 

雄蜂

雄蜂(雄性蜜蜂)是蜂群中最大的蜜蜂。它们通常只在春末和夏季出现。雄蜂的头部比蚁后或工蚁的头部都要大得多,它的复眼在头顶相遇。雄蜂没有毒刺、花粉篮或蜡腺。它们的主要功能是在交配飞行期间使处女蜂后受精。雄蜂在出现约一周后性成熟,并在交配后立即死亡。尽管雄蜂对蜂巢没有任何有用的工作,但它们的存在被认为对正常的蜂群功能很重要。

虽然雄蜂通常依靠工蜂获取食物,但它们可以在 4 天后在蜂巢内自行进食。由于雄蜂吃的食物是工蜂的三倍,过多的雄蜂可能会给蜂群的食物供应带来额外的压力。雄蜂在蜂巢中待到大约 8 天大,之后它们开始进行定向飞行。从蜂巢起飞通常发生在中午和下午 4:00 之间。从未观察到雄蜂从鲜花中获取食物。

当秋天开始寒冷的天气并且花粉/花蜜资源变得稀缺时,雄蜂通常被迫离开蜂巢并饿死。然而,无皇后的蜂群会允许它们无限期地留在蜂巢中。

 

工蜂

工蜂是最小的,占蜂群的大多数。它们是性未发育的雌性,在正常的蜂巢条件下不产卵。工蜂具有专门的结构,如育雏食物腺、气味腺、蜡腺和花粉篮,使他们能够完成蜂巢的所有工作。在它们成年后的最初几周,他们清洁和擦亮蜂房、喂养育雏、照顾蜂后、清除碎屑、处理传入的花蜜、建造蜂蜡梳、守卫入口、空调和通风。后来作为田间蜜蜂,它们觅食花蜜、花粉、水和蜂胶(植物汁液)。

夏季工人的寿命约为6周。秋季饲养的工蚁可存活长达 6 个月,使群体能够在冬季存活,并在春季死亡前协助饲养新的一代。

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Honey bees are social insects, which means that they live together in large, well-organized family groups. Social insects are highly evolved insects that engage in a variety of complex tasks not practiced by the multitude of solitary insects. Communication, complex nest construction, environmental control, defense, and division of the labor are just some of the behaviors that honey bees have developed to exist successfully in social colonies. These fascinating behaviors make social insects in general, and honey bees in particular, among the most fascinating creatures on earth.

A honey bee colony typically consists of three kinds of adult bees: workers, drones, and a queen. Several thousand worker bees cooperate in nest building, food collection, and brood rearing. Each member has a definite task to perform, related to its adult age. But surviving and reproducing take the combined efforts of the entire colony. Individual bees (workers, drones, and queens) cannot survive without the support of the colony.

In addition to thousands of worker adults, a colony normally has a single queen and several hundred drones during late spring and summer (Figure 1). The social structure of the colony is maintained by the presence of the queen and workers and depends on an effective system of communication. The distribution of chemical pheromones among members and communicative “dances” are responsible for controlling the activities necessary for colony survival. Labor activities among worker bees depend primarily on the age of the bee but vary with the needs of the colony. Reproduction and colony strength depend on the queen, the quantity of food stores, and the size of the worker force. As the size of the colony increases up to a maximum of about 60,000 workers, so does the efficiency of the colony.

Queen

Each colony has only one queen, except during and a varying period following swarming preparations or supersedure. Because she is the only sexually developed female, her primary function is reproduction. She produces both fertilized and unfertilized eggs. Queens lay the greatest number of eggs in the spring and early summer. During peak production, queens may lay up to 1,500 eggs per day. They gradually cease laying eggs in early October and produce few or no eggs until early next spring (January). One queen may produce up to 250,000 eggs per year and possibly more than a million in her lifetime.

A queen is easily distinguished from other members of the colony. Her body is normally much longer than either the drone’s or worker’s, especially during the egg-laying period when her abdomen is greatly elongated. Her wings cover only about two-thirds of the abdomen, whereas the wings of both workers and drones nearly reach the tip of the abdomen when folded. A queen’s thorax is slightly larger than that of a worker, and she has neither pollen baskets nor functional wax glands. Her stinger is curved and longer than that of the worker, but it has fewer and shorter barbs. The queen can live for several years—sometimes for as long as 5, but average productive life span is 2 to 3 years.

The second major function of a queen is producing pheromones that serve as a social “glue” unifying and helping to give individual identity to a bee colony. One major pheromone—termed queen substance—is produced by her mandibular glands, but others are also important. The qualities of the colony depend largely on the egg-laying and chemical production capabilities of the queen. Her genetic makeup—along with that of the drones she has mated with—contributes significantly to the quality, size, and temperament of the colony.

About one week after emerging from a queen cell, the queen leaves the hive to mate with several drones in flight. Because she must fly some distance from her colony to mate (nature’s way of avoiding inbreeding), she first circles the hive to orient herself to its location. She leaves the hive by herself and is gone approximately 13 minutes. The queen mates, usually in the afternoon, with seven to fifteen drones at an altitude above 20 feet. Drones are able to find and recognize the queen by her chemical odor (pheromone). If bad weather delays the queen’s mating flight for more than 20 days, she loses the ability to mate and will only be able to lay unfertilized eggs, which result in drones.

After mating the queen returns to the hive and begins laying eggs in about 48 hours. She releases several sperm from the spermatheca each time she lays an egg destined to become either a worker or queen. If her egg is laid in a larger drone-sized cell, she does not release sperm. The queen is constantly attended and fed royal jelly by the colony’s worker bees. The number of eggs the queen lays depends on the amount of food she receives and the size of the worker force capable of preparing beeswax cells for her eggs and caring for the larva that will hatch from the eggs in 3 days. When the queen substance secreted by the queen is no longer adequate, the workers prepare to replace (supersede) her. The old queen and her new daughter may both be present in the hive for some time following supersedure.

New (virgin) queens develop from fertilized eggs or from young worker larvae not more than 3 days old. New queens are raised under three different circumstances: emergency, supersedure, or swarming. When an old queen is accidentally killed, lost, or removed, the bees select younger worker larvae to produce emergency queens. These queens are raised in worker cells modified to hang vertically on the comb surface (Figure 2). When an older queen begins to fail (decreased production of queen substance), the colony prepares to raise a new queen. Queens produced as a result of supersedure are usually better than emergency queens since they receive larger quantities of food (royal jelly) during development. Like emergency queen cells, supersedure queen cells typically are raised on the comb surface. In comparison, queen cells produced in preparation for swarming are found along the bottom margins of the frames or in gaps in the beeswax combs within the brood area.

Drones

Drones (male bees) are the largest bees in the colony. They are generally present only during late spring and summer. The drone’s head is much larger than that of either the queen or worker, and its compound eyes meet at the top of its head. Drones have no stinger, pollen baskets, or wax glands. Their main function is to fertilize the virgin queen during her mating flight. Drones become sexually mature about a week after emerging and die instantly upon mating. Although drones perform no useful work for the hive, their presence is believed to be important for normal colony functioning.

While drones normally rely on workers for food, they can feed themselves within the hive after they are 4 days old. Since drones eat three times as much food as workers, an excessive number of drones may place an added stress on the colony’s food supply. Drones stay in the hive until they are about 8 days old, after which they begin to take orientation flights. Flight from the hive normally occurs between noon and 4:00 p.m. Drones have never been observed taking food from flowers.

When cold weather begins in the fall and pollen/nectar resources become scarce, drones usually are forced out into the cold and left to starve. Queenless colonies, however, allow them to stay in the hive indefinitely.

Workers

Workers are the smallest and constitute the majority of bees occupying the colony. They are sexually undeveloped females and under normal hive conditions do not lay eggs. Workers have specialized structures, such as brood food glands, scent glands, wax glands, and pollen baskets, which allow them to perform all the labors of the hive. They clean and polish the cells, feed the brood, care for the queen, remove debris, handle incoming nectar, build beeswax combs, guard the entrance, and air-condition and ventilate the hive during their initial few weeks as adults. Later as field bees they forage for nectar, pollen, water, and propolis (plant sap).

The life span of the worker during summer is about 6 weeks. Workers reared in the fall may live as long as 6 months, allowing the colony to survive the winter and assisting in the rearing of new generations in the spring before they die.