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陆向明 · 发表于 2023-12-10 14:05:06

本帖最后由陆向明于 2023-12-10 14:38 编辑

Queen Pheromone(S)
蜂王信息素

We humans hear the term “queen” and immediately project all the royal attributes of leadership and omnipotence upon the mother of the colony.
我们人类见到“蜂王”这个字眼，立刻会把国王的象征--至高无上的地位和无所不能的权利--捧给这位蜂群的母亲。

But is this really the case?
果真如此吗？

Does the queen bee really “rule” the colony, with the sterile workers acting as subordinate subjects?
蜂王真的统治着蜂群，让不育的工蜂充当奴隶吗？

The feeling that I get is that bee society is not that way at all.
我感觉蜜蜂社会全然不是那样的。

It appears to me that the superorganism—the colony—is the true decision maker, and that the queen functions merely as the ovary and reservoir of genetics for the colony, and that her pheromones function mainly as reports to the superorganism as to the status of the ovary.
在我看来，超个体--蜂群--才是真正的决策者，而蜂王的作用仅仅是作为蜂群的卵巢和遗传库，她的信息素的功能主要是向超个体报告卵巢的状态。

Indeed, Seeley (1995) describes “messenger bees that pick up the queen’s pheromones and then travel about the broodnest actively dispersing this olfactory indicator of the queen’s presence.”
事实上，Seeley（1995）描述了“信使蜂拾取蜂王信息素，活跃在巢房的每一个角落，传播着这些表示蜂王存在的嗅觉物质。”

As long as the “ovary” is properly mated and functioning well, the colony is content—the presence of both brood pheromone and queen pheromone suppresses ovary development in the workers.
只要“卵巢”交配恰当并且功能良好，蜂群就会感到满足--幼虫信息素和蜂王信息素的存在都会抑制工蜂的卵巢发育。

However, should the “healthy ovary” signals falter or disappear (Fig. 1), the colony will immediately swing into action to replace the queen/ovary without hesitation or sentiment, via supersedure, or the initiation of emergency queen cells.
然而，如果“健康卵巢”信号衰退或消失（图1），蜂群将立即毫不犹豫地采取行动，通过交替或急造王台更换蜂王。

I’m using the term “queen pheromone” (QP) generically, since the queen produces a bouquet of pheromonal components from different glands, with multiple functions.
我通常使用术语“蜂王信息素”（QP），因为蜂王从不同的腺体产生一组多种功能的信息素组分。

Current terminology has moved from “queen mandibular pheromone” to “queen retinue pheromone,” since not all components necessary for retinue formation are produced by her mandibular glands.
目前的术语已经从“蜂王下颌信息素”转变为“蜂王伴生信息素”，因为并非所有信息素的构成组分都由她的下颌腺产生。

So far, nine components of the full retinue pheromone have been named, but it is clear that there are still others yet to be identified.
到目前为止，完整的伴生信息素的九个成分已被命名，但显然还有其他成分尚未确定。

This brings up a fascinating aspect of QP—there is a genetic component involved in worker recognition of the individual components.
这带来了 QP 的一个令人着迷的方面——工蜂的参与识别各组分的基因组。

Some strains of bees do not recognize the queen if certain components are not present in the right amount!
如果某些组分的含量不正确，某些蜜蜂群体就无法识别蜂王！

Practical app: This finding makes me wonder if that is why it is difficult to introduce queens of some strains into unrelated colonies, and whether this might be related to the substantial amount of rapid supersedures sometimes observed after introducing purchased queens. If you are introducing queens of a different stock than the recipient colony, the workers simply may not recognize her pheromonal signals as being “right”!
实践应用：这一发现让我想知道这是否就是为什么很难将某些群体的蜂王介入不相关的蜂群的原因，以及这是否可能与介入购买的蜂王后有时观察到的大量快速取代有关。如果您引入的蜂王与接收群体的种群不同，工蜂可能根本无法识别出她的信息素信号是“正确的”！

Also of interest is that the queen apparently signals to the colony as to how well she was mated—queens that are inseminated by multiple drones produce QP that is more attractive to workers than that produced by queens mated to a single drone (Richard 2007). Kocher (2009) found “that the queen pheromone blend is modulated by the reproductive status of the queens, and workers can detect these subtle differences and are more responsive to queens with higher reproductive potential.” The quality of the QP produced by any individual queen may have considerable impact upon a colony, regardless of her genetics—Grozinger (2003) found that QP affected the expression of several hundred genes in workers!
同样有趣的是，蜂王显然向蜂群发出信号，表明她交配的多么好--由多只雄蜂授精的蜂王产生的QP比与单个雄蜂交配的蜂王产生的QP对工蜂更具吸引力 (Richard 2007)。Kocher（2009）发现“蜂王信息素是由蜂王的繁殖状态调节的，工蜂可以感受到这些细微的差异，并对具有更高繁殖潜力的蜂王更敏感。”任何蜂王产生的 QP 质量都可能对蜂群产生不可忽视的影响，运用基因分析——Grozinger (2003) 发现 QP 影响工蜂中数百个基因的表达！

Practical app: Poorly-mated queens (such as those forced to take mating flights during cool or rainy weather) may be quickly superseded. A well-mated queen, on the other hand, functions as a pheromonal “cheerleader” to energize and invigorate the colony, and to boost morale.
实践应用：交配糟糕的蜂王（例如那些被迫在低温或下雨天气婚飞的蜂王）可能很快被交替。相反，交配好的蜂王则充当着信息素的“啦啦队长”，激励和振奋着团队，使蜂群士气高涨。

Colony Reproduction

蜂群繁殖

There are two ways for a colony to disseminateits genetics (and epigenetics—we’ll get to this later) into future generations:by either the production of drones, or by division of the colony by swarming(an unusual process, in that the “parent” leaves, and the “offspring” staysbehind). As you can see in Figure 1, the primer pheromones, pollen income, andvitellogenin are all involved in the “decision making” of colony reproduction.

一个蜂群可以通过两种方式将其基因（以及表观遗传学——我们稍后会谈到）传递给后代：要么通过雄峰，要么通过分蜂（这是一个不寻常的过程，因为“父母”离开，“后代”留下）。如图 1 所示，基本的信息素、花粉收入和卵黄蛋白原都影响蜂群繁殖的“决策”。

待续。。。。。。。。

陆向明 · 发表于 2023-12-10 15:52:28

Drone Production
雄峰生产

Compared to the risky and expensive enterprise of throwing off a swarm, the creation of expendable drones is a cheap gamble for a colony at getting a portion of its genes into the next generation.
为了将基因传递给下一代，与分蜂的风险和代价相比，生产雄峰对于蜂群说是一场廉价的冒险。

Even though any particular drone has little chance of actually mating, the colony’s investment in that drone is minimal.
尽管任何特定的雄蜂实际交配的机会都很小，但蜂群对该雄蜂的投入也很少。

But by producing thousands of drones, the colony has a good chance at disseminating its genes when virgin queens from other colonies fly out to mate.
但是，通过生产成千只雄蜂，当来自其他蜂群的处女王飞出去交配时，该蜂群就有很好的机会传播其基因。

Each drone takes a certain amount of colony resources to produce and feed, so the hive carefully regulates the amount and timing of drone production.
每只雄蜂都需要一定数量的蜂群资源来生产和喂养，因此蜂群会周密调节雄蜂生产的数量和时间。

The mechanisms for such regulation are not completely understood (reviewed recently by Boes 2010).
这种调节的机制尚未完全被破解（Boes 2010 最近回顾）。

Since the only purpose of drones is to mate with flying queens (generally from other colonies), it is obvious that the best time to rear drones is when they actually have a chance at “getting lucky”—during swarming season, when the air is full of lusty young virgins.
由于雄蜂的唯一目的是与飞翔的蜂王（通常来自其他群体）交配，显然，培育雄蜂的最佳时机是它们真正有机会“走运”的时候——空中满是精力充沛的年轻处女的分蜂季节。

At other times of the year, the only value of maintaining a population of drones is that one may have a chance encounter with a supersedure or emergency queen.
在一年中的其他时间，维持雄蜂种群的唯一价值是，可能有机会遇到交替蜂王或急造蜂王。

Drones are produced, as a rule, only once a colony has reached substantial size, and then only when there is abundant pollen coming into the hive, the presence of which unleashes a cascade of pheromonal signals, resulting in a vitellogenin-rich population of nurse bees awash in jelly, and thus kicking off feedback loops of queen and brood pheromone production.
通常，只有当蜂群达到相当大的规模，并且只有当有大量花粉进入蜂巢时，才会生产雄蜂，花粉的存在会释放一系列信息素信号，从而产生富含卵黄蛋白原的能大量分泌王浆的哺育蜂群体，从而启动了蜂王信息素和幼虫信息素产生的反馈循环。

When colony nutrition is poor, the freeloading drones are summarily evicted.
当蜂群营养不良时，不劳而获的雄蜂就会被立即驱逐。

Practical apps: the presence of drone brood is a great indicator that a colony is in good nutritional shape; conversely, their eviction might suggest the need for supplemental feeding.
实践应用：雄蜂巢的存在是一个很好的指标，表明蜂群营养状况良好；相反，他们被驱逐可能表明需要补充喂养。

Steve Taber demonstrated that one could get colonies to produce drones even during winter by feeding them pollen patties.
史蒂夫•泰伯（Steve Taber）证明，即使在冬天，通过喂食花粉饼，也可以让蜂群生产出雄峰。

Colonies will utilize introduced drone comb to rear more drones than they would without such added comb, but at an energetic cost to the colony (Seeley 2002).
与没有引入额外的蜂巢相比，蜂群将利用引入的雄峰蜂巢来饲养更多的雄峰，但这会给蜂群带来能量成本（Seeley 2002）。

Don’t worry about swarming at least until after a colony has produced drones—as far as I can tell, a colony invariably initiates drone production prior to building swarm cells.
至少在蜂群生产出雄峰之前不要担心蜂群——据我所知，蜂群总是在分蜂之前开始生产无雄峰。

This makes evolutionary sense, since in bad times any rare surviving colonies would want to ensure that there were some drones present to inseminate the new queen (surprisingly, apparently no one has tested to see whether the presence of drones is essential for swarming).
这在进化上是有道理的，因为在困难时期，任何稀有的幸存群体都会希望确保有一些雄蜂
来为新蜂王授精（令人惊讶的是，显然没有人测试过雄蜂的存在对于分蜂是否是必要的）。

Practical Tip: During pollen dearths, colonies stop rearing drones. At that time, varroa are forced into worker brood (this generally occurs during August and September). The combination of nutritional stress and high varroa infestation often leads to virus epidemics in the workers that can spell death for the colony (Sumpter and Martin 2004). Thus, this is the most critical time to monitor mite levels.
实用提示：在花粉缺乏期间，蜂群会停止饲养雄蜂。那时，瓦螨被迫进入工蜂窝（这通常发生在八月和九月）。营养压力和高瓦螨感染的结合常常导致工蜂中病毒流行，从而导致蜂群灭亡（Sumpter 和 Martin 2004）。因此，这是监测螨虫水平的最关键时期。

陆向明 · 发表于 2023-12-11 08:00:50

Swarming
分蜂

Swarm production, as opposed to drone rearing, is a risky large-scale investment, with limited potential of founding a successful new colony (most swarms in nature do not survive the winter).
与雄峰生产相比，分蜂生产是一项有风险的大规模投资，建立成功的新蜂群的潜力有限（自然界中的大多数蜂群都无法过冬）。

The tradeoff is that a swarm has a much better chance at passing on its genes (all of a colony’s genetics are carried in a prime swarm –leaving a related daughter in the hive to carry on, with the infusion of genes from drones of other successful nearby colonies).
有利的是，蜂群有更好的机会传递其基因（蜂群的所有遗传都在原蜂群中携带，将其他成功蜂群的基因注入到蜂群中，将相关的子代留在蜂巢中继续繁衍。附近的殖民地）。

Swarming behavior is complex.
分蜂行为很复杂。

It is strongly seasonal, but appears to be more directly due to colony population size (dilution of QP), the nurse to brood ratio (Vg and BP), pollen availability and the resultant abundance of jelly and Vg (Pridal and Sustek 2000), and age of the queen (QP). (I’m curious as to the ratio of open to sealed brood—in an expanding colony, there is a great deal of larvae secreting BP, but in a pre-swarming colony, most of the brood is sealed).
它具有很强的季节性，但似乎更直接地归因于蜂群规模（QP 稀释）、育雏保育员的比率（Vg 和 BP）、花粉可用性以及由此产生的王浆和 Vg 丰度（Pridal 和 Sustek 2000），和女王的年龄（QP）。（我很好奇未封盖与封盖幼虫的比例——在一个正在扩张的群体中，有大量的幼虫分泌BP，但在一个预分蜂的群体中，大部分幼虫是封盖的）。

Lenskya (1981) observed that in crowded hives, the queen avoided the lower edges of the brood combs, and found that by applying a combination of QP and queen “footprint” pheromone to the lower edges of combs, that he could inhibit the bees from building queen cells there.
Lenskya (1981) 观察了拥挤的蜂巢，蜂王如何使蜂巢的下边缘无效：发现通过在蜂巢的下边缘应用 QP 和蜂王“足迹”信息的组合，可以抑制蜜蜂在那里建造王台。

Because of the substantial economic significance of swarming to the beekeeper, management for swarm prevention has been vigorously debated, but I’ll leave that discussion for another article!
由于分蜂对养蜂人具有重大的经济意义，阻止分蜂的管理一直存在激烈的争论，我将把这个讨论留到另一篇文章！

Some exceptionally large swarm cells from a strong colony that has been enjoying an abundance of forage.
一些异常大的分出群来自一个强大的蜂群，该蜂群一直在享受丰富的饲料。

The queen, shortly before she is ready to emerge as an adult, emits a pheromonal signal for the workers to remove the cell wax from the tip of her silk cocoon (cell at left).
蜂王在即将出台之前，会向工蜂发出信息素信号，以清除其丝茧尖端的蜡（王台头部）。

Photo by the author.
作者拍的照片

Practical app: In short, keep these facts in mind: a colony tends to swarm if it has an aging queen, is in the best nutritional shape, has an abundance of stores (has filled the combs), is crowding the cavity with bees, and the queen has filled every available cell with brood. You can minimize swarming by manipulating the hive to change any of these factors during the relatively short seasonal swarm window.
实践应用：简而言之，请记住这些事实：如果一个蜂群有一个年老的蜂王、处于最佳的营养状态、有丰富的储备（已经填满了蜂巢）、蜂巢里挤满了蜜蜂，蜂王已经把所有可用的巢房都装满了幼虫，那么它就会倾向于分蜂。您可以通过在相对较短的季节性蜂群窗口期间操纵蜂巢来改变这些因素，从而最大限度地减少分蜂。

陆向明 · 发表于 2023-12-11 08:29:48

Survival Mode
生存模式

The colony will go into survival mode during periods of nutritional duress.
在食物匮乏期间，蜂群将进入生存模式。

The pheromonal trigger for the production of long-lived “winter” or diutinus bees appears to be the presence of QP concurrent with the absence of BP, causing bees that would normally be nurses to store Vg in their fat bodies (as opposed to converting it to jelly) in order to wait out the dearth.
信息素触发生产长寿“越冬”蜂，蜜蜂的QP 的存在与 BP 的缺乏同时发生，导致通常作为哺育的蜜蜂将Vg储存在其脂肪体内（而不是转化为王浆）以等待饥饿结束。

陆向明 · 发表于 2023-12-11 08:35:25

Other Factors
其他因素

I would be remiss not to mention that there is more to the allocation of the workforce than just the primer pheromones in Figure 1.
如果我不提及劳动力的分配，仅仅提及图 1 中的信息素，那就太失职了。

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