萤火虫的研究、保护及开发利用进展

萤火虫是萤科和凹眼萤科昆虫的统称,既是科学研究的良好材料,也具有巨大的产业潜力。本文从分类学(包括分类现状和分子系统学研究进展)、生物学(包括生活史、分布、天敌、视蛋白、人工光照的影响、生物指示作用、体内共生菌等)、行为学(包括发光、求偶、交配、取食、防御、报警、拟态、叠背、成虫后多次蜕皮等)等方面全面梳理了国内外萤火虫基础研究的进展,分析了萤火虫的消亡原因(环境污染、旅游影响、科普欠缺、生物入侵及其它等)、保护情况并提出了保护萤火虫的若干措施建议,系统地阐述了当前国内外萤火虫开发利用的现状,最后从萤火虫的科学研究、资源保护、人工养殖、产业发展等方面提出了建议,以期推动萤火虫的研究和资源开发工作。     

萤火虫的发光行为及其功能起源

发光生物因其活体能自发荧光而倍受关注,萤火虫(鞘翅目:萤科)是被研究最多的发光生物,因为成虫发光在其性选择以及求偶交配中起着重要作用.由于萤火虫的发光是一个消耗能量(ATP),的化学反应过程,因而其发光在成虫之外的虫态也具有重要意义.本文就萤火虫成虫和幼虫的发光行为、功能意义进行了综述,并结合萤火虫的饲养观察对其卵和蛹的发光行为进行了描述,探讨了卵和蛹的形态阶段发光的生物学功能以及生物荧光的起源进化.这将有助于理解萤火虫及其它生物自发荧光的本质和起源进化过程.     

萤火而今,飞破秋夕

正日本有一部著名的动画电影,名叫《萤火虫之墓》,是以萤火虫微弱的生命力作为意象命名的。自古以来,中国就是亚洲文化的中心,萤火虫意象的诸多文化特点,也基本来源于中国传统文化。其实,从古人对萤火虫的命名就能看出来,萤火虫注定要成为一种独特的昆虫。唐代用于辅助诸皇子学习的类书《初学记》记载了萤火虫的别名,分别是耀夜、夜光、宵燭、景天、熠燿、丹     

萤火虫发光的化学机制

在若干昆虫中,普遍存在着生物冷光(Bioluminescence)现象。这些发光的昆虫大多数属于鞘翅目、萤科,通常称做萤火虫。这些昆虫发光有专门的复器,也是一种交配的信号。关于萤火虫发光的化学机理问题,不少学者进行过研究探讨。1947年,McElroy最早用北美洲萤火虫photinus pyralis作材料进行研究,发现萤火虫生物冷光要求ATP。1952年,B.L.斯特勒和J.R.托特尔用萤火虫生物冷光探测ATP,其灵敏度可达10~(-15)M(ATP);同年,Harvey研究发现,萤火虫发冷光的颜色是不同的,这是因为不同种的萤火虫含有不同类型的萤光素/萤光素酶系统。以后,M.德卢卡又进行了一系列的研究,提出了萤火虫发光反应的现代知识,这个被概述在图1中。     

萤火虫(鞘翅目:萤科)两性交流中的闪光信号

对国内外萤火虫两性交流闪光信号的研究进行了综述,萤火虫发光器因种而异,多数发出黄绿色萤光,闪光信号的频率、光谱、强度及其时空分布的闪光模式包含着两性交流信息。萤火虫闪光交流系统有两种分类方法,其一是萤火虫具两个类型的闪光信号交流系统,及系统和系统,前者多在旧大陆,后者多在新大陆;其二是萤火虫具6个类型闪光信号交流系统,即HP,LL,LC,PR,CR和LB型,其中PR型与系统相对应,HP型与系统对应。萤火虫两性交流闪光信号常因时间和空间上的差异及外界物体的干扰使两性闪光交流的效率受到影响。萤火虫两性交流的闪光信号起源于鞘翅目的幼虫阶段,并起警戒天敌的作用,经过两性选择成为成虫两性交流的一种途径,进而成为新大陆的一些萤火虫间捕食猎物和逃避天敌的生存策略。     

萤火虫的奥秘

近年来,科学家们不仅揭开了萤火虫发光的秘密,而且发现了这种昆虫在生活过程中的一些奇特的现象。从卵到成虫萤火虫(一种甲虫)一开始以卵的形式藏在土里,很多种萤火虫的卵就已经能显现出极其微弱的亮光,就象那希望的闪     

中国萤火虫云南窗萤荧光素酶cDNA的克隆表达和序列分析(英文）

从一种来自中国日行性萤火虫（云南窗萤）发光器官mRNA中克隆、测序并表达了有功能的荧光素酶。云南窗萤荧光素酶的cDNA序列有1 647个碱基,编码548个氨基酸残基。从推测得到的氨基酸序列的比对分析得出：云南窗萤的荧光素酶与来自Lampyris noctiluca, L. turkestanicus和Nyctophila cf. caucasica三种萤火虫的荧光素酶有97.8%的序列一致性。从推测得出的氨基酸序列进行系统发育分析,其结果表明：云南窗萤和Lampyris+Nyctophila聚在一起, 与同属的发光强夜行性的萤火虫不形成的单系。云南窗萤荧光素酶在大肠杆菌中表达的条带大约70 kDa,并且在有荧光素存在时发出黄绿色荧光。对荧光素酶的结构模拟和分析表明,云南窗萤荧光素酶基因的氨基端和羧基端结构域之间的裂沟处存在这5个多肽环,这正是从其他荧光素酶推测得到的催化荧光反应时的底物结合位点。云南窗萤和窗萤属的其他3种萤火虫的荧光素酶相比,有13个不同氨基酸位点,位于模拟分子结构的表面。对于这些多肽环、不同氨基酸残基和晶体结构的进一步研究有利于解释日行和夜行性萤火虫荧光素酶的差异。     

云南窗萤的形态学及其生物学特性（鞘翅目：萤科）

研究了云南窗萤卵到成虫不同发育阶段的形态学。根据野外观察和实验室饲养结果,记录了其生物学特性。卵、幼虫、蛹和雌成虫可发弱光,但雄成虫仅在受刺激时才发出更弱的光,云南窗萤可归属于昼行性萤火虫,而化学信号则是其雌雄在求偶时的主要识别方式。     

中国萤火虫云南窗萤荧光素酶cDNA的克隆表达和序列分析

从一种来自中国日行性萤火虫(云南窗萤)发光器官mRNA中克隆、测序并表达了有功能的荧光素酶.云南窗萤荧光素酶的cDNA序列有1647个碱基,编码548个氨基酸残基.从推测得到的氨基酸序列的比对分析得出:云南窗萤的荧光素酶与来自Lampyris noctiluca,L.turkestanicus和Nyctophila cf.caucasica三种萤火虫的荧光素酶有97.8%的序列一致性.从推测得出的氨基酸序列进行系统发育分析,其结果表明:云南窗萤和Lampyris Nyctophila聚在一起,与同属的发光强夜行性的萤火虫不形成的单系.云南窗萤荧光素酶在大肠杆菌中表达的条带大约70kDa,并且在有荧光素存在时发出黄绿色荧光.对荧光素酶的结构模拟和分析表明,云南窗萤荧光素酶基因的氨基端和羧基端结构域之间的裂沟处存在这5个多肽环,这正是从其他荧光素酶推测得到的催化荧光反应时的底物结合位点.云南窗萤和窗萤属的其他3种萤火虫的荧光素酶卡目比,有13个不同氨基酸位点,位于模拟分子结构的表面.对于这些多肽环、不刚氨基酸残基和晶体结构的进一步研究有利于解释日行和夜行性萤火虫荧光素酶的差异.     

小小萤火虫，我们怎能失去你!

夏夜纳凉时观赏小小萤火虫闪烁微光,是许多成人在童年时期的美好记忆。然而现在还有多少孩子亲眼看过萤火虫,发出惊喜的尖叫？2008年诺贝尔化学奖授予了生物发光领域的三位美国专家,萤火虫的科技价值在中国还一直并没被重视;作为公认的环境指示物种,徘徊在灭亡边缘的萤火虫还预示着环境危机和生态退化……     

Spontaneous flash communication of females in an Asian firefly

Fireflies are well known for the use of bioluminescence for sexual communication. In species using flash signals for pair formation, species and sexual identity are conferred by flash timing parameters such as flash duration, flash interval, flash number, and response delay. In dialog fireflies in North America, the male is the advertiser and the female is the responder. In these species, the male flash signal parameter varies depending on species, but the female flash signal parameter is limited only to response delay. However, in fireflies other than dialog fireflies, sexual flash communication is not well studied. Although many female-advertisement-like fireflies are reported, we have no confirmed case of sexual communication in a female-advertisement species. Here, we report the sexual flash communication of an Asian firefly, Luciola (Hotaria) parvula, in which the female flashes spontaneously. By using an electronic firefly, we confirm experimentally that males are specifically attracted to flashes with a female-specific flash duration. This is the first experimental report of sexual communication of a female advertiser in firefly communication. In this species, females call males usually with spontaneous flashes unlike dialog fireflies.     

Variation in flash speed of Japanese firefly,Luciola cruciata (Coleoptera: Lampyridae), identifies distinct southern “quick‐flash” population on Goto Islands,Japan

Luciola cruciata, a bioluminescent firefly endemic to Japan, communicates with species‐specific flashing patterns. Previous studies reported that flying males possess three distinct flashing frequencies: slow (4 s), intermediate (3 s) and fast (2 s). Our study focused on geographical variations in flashing patterns in the northwestern part of the Kyushu region of southern Japan (including its isolated islands) and their relationship to the mitochondrial COII region of firefly DNA. We found that “quick‐flash (1 s)” fireflies occur in the Goto Islands, at the western tip of Kyushu, and that, although they possess a unique haplotype, they do not differ much genetically from “fast‐flash” fireflies inhabiting mainland Kyushu. In contrast, although fireflies inhabiting the Tsushima and Iki Islands also possess a unique haplotype, their flashing patterns were the same (fast‐flash) as fireflies inhabiting mainland Kyushu. We determined that flash patterns of L. cruciata in Japan should be classified according to four flashing patterns: slow, intermediate, fast and quick.     

Adult firefly abundance is linked to weather during the larval stage in the previous year

1. Much is known about the brief adult phase of fireflies. However, fireflies spend a relatively long developmental period under the soil surface. Climatic and soil conditions may directly affect the eggs, larvae, and pupae, and also affect them indirectly through predators, competitors, and prey items. Climatic conditions during the early life stages of this iconic species are therefore relevant to their hypothesised decline within the context of global warming. 2. Data on the abundance of fireflies were extracted from the publicly available citizen dataset across North America over a period of 9 years. The effects of weather in the 24 months before the observations of firefly abundance were documented based on 6761 observations. 3. Climatic conditions during both the larval and adult phases have a non‐linear effect on adult firefly abundance. Maximum winter and spring temperatures and mean precipitation in the 20‐month period before the observations had the greatest impact on the abundance of firefly adults. Low maximum soil moisture during the 5–19 months preceding the observations affected the adult abundance negatively, and high maximum soil moisture affected it positively. 4. After correcting the firefly abundance for these weather effects, it is estimated that the abundance of fireflies increased over the time period of this study. 5. This study suggests that early life climatic conditions have a small but significant impact on adult firefly abundance with a total R² of 0.017.     

Examining the Role of Cuticular Hydrocarbons in Firefly Species Recognition

During animal courtship, multiple signals transmitted in different sensory modalities may be used to recognize potential mates. In fireflies (Coleoptera: Lampyridae), nocturnally active species rely on long-range bioluminescent signals for species, sex, and mate recognition, while several diurnally active species rely on pheromonal signals. Although in many insects non-volatile cuticular hydrocarbons (CHC) also function in species and sex discrimination, little is known about the potential role of CHC in fireflies. Here, we used gas chromatography to characterize species and sex differences in the CHC profiles of several North American fireflies, including three nocturnal and two diurnal species. Additionally, we conducted behavioral bioassays to determine whether firefly males (the searching sex) were differentially attracted to extracts from conspecific vs. heterospecific females. Gas chromatography revealed that nocturnal Photinus fireflies had low or undetectable CHC levels in both sexes, while diurnal fireflies showed higher CHC levels. No major sex differences in CHC profiles were observed for any firefly species. Behavioral bioassays demonstrated that males of the diurnal firefly Ellychnia corrusca were preferentially attracted to chemical extracts from conspecific vs. heterospecific females, while males of the remaining species showed no discrimination. These results suggest that while CHC may function as species recognition signals for some diurnal fireflies, these compounds are unlikely to be important contact signals in nocturnal Photinus fireflies.     

A New Type of Synchronized Flashing in a North American Firefly

Continuous synchronic flashing occurs in stationary and flying male congregations of members of some firefly species in Southeast Asia. In the present paper, low-light videography and photometry was used to demonstrate that synchrony occurs in the North American genus Photuris. We found that the Georgia coastal plain firefly Ph. frontalis flashed synchronically. From a distance, the synchronic flashes of a population of flying Ph. frontalis appeared to occur in a continuous synchrony. However, when pairs of males were viewed, it was difficult to verify that their flashing was continuously synchronic. In the laboratory, caged fireflies flashed synchronically, stopped, and then flashed synchronically again. To study this flash behavior, recordings of individual and group flashing were analyzed statistically to validate the conclusions about rhythm and synchrony. Although the mass synchrony appeared continuous, the individual flying males switch on and off, coming in again on the beat. The synchrony in Ph. frontalis is common and pervasive rather than rare and sporadic, as shown by other North American fireflies. The precision of the frontalis synchrony approaches that of Southeast Asian fireflies. The intermittent synchrony of this firefly reinforces the evidence that there is a diversity of synchronies in fireflies.     

A spiking neuron model for synchronous flashing of fireflies

Certain species of fireflies show a group behavior of synchronous flashing. Their synchronized and rhythmic flashing has received much attention among many researchers, and there has been a study of biological models for their entrainment of flashing. The synchronous behavior of fireflies resembles the firing synchrony of integrate-and-fire neurons with excitatory or inhibitory connections. This paper shows an analysis of spiking neurons specialized for a firefly flashing model, and provides simulation results of multiple neurons with various transmission delays and coupling strengths. It also explains flashing patterns of some firefly species and examines the synchrony conditions depending on transmission delays and coupling strengths.     

Firefly Femmes Fatales: A Case Study in the Semiotics of Deception

Mimicry and deception are two important issues in studies about animal communication. The reliability of animal signs and the problem of the benefits of deceiving in sign exchanges are interesting topics in the evolution of communication. In this paper, we intend to contribute to an understanding of deception by studying the case of aggressive signal mimicry in fireflies, investigated by James Lloyd. Firefly femmes fatales are specialized in mimicking the mating signals of other species of fireflies with the purpose of attracting responding males to become their prey. These aggressive mimics are a major factor in the survival and reproduction of both prey and predator. It is a case of deception through active falsification of information that leads to efficient predation by femmes fatales fireflies and triggered evolutionary processes in their preys’ communicative behaviors. There are even nested coevolutionary interactions between these fireflies, leading to a remarkable system of deceptive and counterdeceptive signaling behaviors. We develop here a semiotic model of firefly deception and also consider ideas advanced by Lloyd about the evolution of communication, acknowledging that deception can be part of the explanation of why communication evolves towards increasing complexity. Increasingly complex sign exchanges between fireflies evolve in an extremely slow pace. Even if deceptive maneuvers are played out time and time again between particular firefly individuals, the evolution of the next level of complexity—and thus the next utterance in the dialogue between species—is likely to take an immense amount of generations.     

Pyrazine emission by a tropical firefly: An example of chemical aposematism?

Although famous for photic courtship displays, fireflies (Coleoptera: Lampyridae) are also notable for emitting strong odors when molested. The identity of volatile emissions and their possible role, along with photic signals, as aposematic warnings of unpalatability have been little explored, especially in tropical species. Pursuant to the observation that the widespread Neotropical fireflies, Photuris trivittata and Bicellonycha amoena, emit pungent odors, glows, and flashes when handled, we investigated their cuticular and headspace chemistry. Gas chromatography–mass spectrometry analyses revealed that both fireflies have species‐specific cuticular hydrocarbon profiles. Photuris trivittata headspace was dominated by 2‐methoxy‐3‐(1‐methylpropyl) pyrazine (hereafter, pyrazine), on the order of 1.59 ng/individual and a suite of sesquiterpenes, while B. amoena emitted 3‐methoxy‐2‐butenoic acid methyl ester and a few ketones. This is the first report of such compounds in fireflies. We investigated the role of pyrazine in P. trivittata's interactions with potential predators: sympatric ants, toads, and bats. Solvent‐washed P. trivittata painted with pyrazine incurred lower ant predation than did their solvent‐washed counterparts. Pyrazine significantly repelled ants at baits in concentrations as low as 9.8 × 10^?4 ng/μl. The toad, Rhinella marina, readily accepted intact fireflies, pyrazine‐coated and uncoated mealworms. Both Myotis nigricans and Molossus molossus bats rejected fireflies, but accepted both pyrazine‐coated and uncoated mealworms. While pyrazine repels ants, its role as an aposematic signal warning other potential predators of firefly distastefulness requires further investigation. Our results underscore the idea that multiple enemies exert conflicting selection on firefly defenses.     

Fireflies with or without prespermatophores: Evolutionary origins and life-history consequences

During mating, some male North American fireflies produce spermatophores from prespermatophores in their paired reproductive accessory glands. Other species of fireflies have neither prespermatophores nor spermatophores. To establish a pattern of spermatophore occurrence across firefly species, we examined the male internal reproductive system in 20 Japanese species belonging to 10 genera for the presence or absence of prespermatophores. Twelve species from seven genera produced prespermatophores, while eight species from three genera did not. Superimposed on a molecular phylogeny of Japanese fireflies based on mitochondrial 16S ribosomal DNA sequences, the basal group was prespermatophore producers. Prespermatophores appear to have been lost in two different lineages. Species without prespermatophores are characterized by degeneration of both the forewings and hindwings, and by body gigantism in females.