An evolutionary treatment of the morphology and physiology of circulatory organs in insects

An overview from an evolutionary perspective is presented on the research of the past 2 decades on insect circulatory organs. Based on various functional morphology it is clear that the flow mode of the dorsal vessel ('heart') has changed during the evolution of hexapods. In all apterygotes and mayflies the flow is bidirectional. In most pterygote insects, however, it is unidirectional. In some endopterygote insects, the direction of the flow alternates. This is achieved by heartbeat reversal, which may have various physiological functions and is a derived condition that probably occurred several times during the course of insect evolution. Special attention is given to the hemolymph flow in body appendages. In ancestral hexapods, they are supplied by arteries, whereas circulation in appendages of higher insects is accomplished by accessory pulsatile organs. These auxiliary hearts are autonomous pumps and exhibit a great diversity in their functional morphology. They represent evolutionary innovations which evolved by recruitment of building blocks from various organ systems and were assembled into new functional units. Almost all pulsatile circulatory organs in insects investigated exhibit a myogenic automatism with a superimposed neuronal control. The neuroanatomy of insect circulatory organs has been investigated only in a small number of species but in considerable detail. Numerous potential peptidergic and a few aminergic mediators could be demonstrated by immunocytochemical and biochemical methods. The cardiotropic effectiveness of these mediators may vary among species and it can be stated that there is no uniform picture of the control of the various circulatory organs in insects. A possible explanation for the differences may lie in the different evolutionary origins of the muscular components. Furthermore, insect circulatory organs may represent important neurohemal releasing sites.     

A Novel Device for the Collection, Storage, Transport, and Delivery of Beneficial Insects, and its Application to Ophraella communa (Coleoptera: Chrysomelidae)

A multi-use device was developed for the collection, short-term storage, transport, and delivery of Ophraella communa (Coleoptera: Chrysomelidae), a biological control agent of common ragweed, Ambrosia artemisiifolia. The device is made from a 125-ml plastic specimen container that can hold O. communa adults or pupae. When used as an aspirator, insect collection and counting times are reduced. O. communa adults and pupae can be stored inside the container at 3°C with median survival of 41 and 21 days, respectively. A cotton wick saturated with water or a 5% sugar solution nourishes insects during transport and the container design minimizes insect mortality by providing an optimum microclimate during insect storage and transport. Designed to protect insects from rainfall and to limit encounters with predators and parasites, the containers can be used for field releases of O. communa adults and pupae. Although the container has been designed and tested for O. communa, it is highly versatile and could possibly be used with a variety of insect species.     

Extracardiac versus cardiac haemocoelic pulsations in pupae of the mealworm (Tenebrio molitor L.)

Pulsations in mechanical pressure of the pupal haemocoele were investigated by means of simultaneous recording from multiple sensors. It has been determined that cardiac and extracardiac haemocoelic pulsations are each regulated by substantially different and quite independent physiological mechanisms. At the beginning and in the middle of the pupal interecdysial period the anterograde heartbeat and extracardiac pulsations occur in similar, but not identical periods. During the advanced pharate adult stage, there appear almost uninterrupted pulsations from different sources: cardiac, extracardiac, intestinal, and the ventral diaphragm.Extracardiac pulsations are associated with pressure peaks of 200-500 Pa, occurring at frequencies of 0.3-0.5 Hz. The effect of heartbeat on haemocoelic pressure is very small, 100- to 500-fold smaller, comprising only some 1 or 2 Pa during the vigorous anterograde systolic contractions. Accordingly, extracardiac pulsations are associated with relatively large abdominal movements from 30-90 μm whereas heartbeat produces movements of only 100-500 nm. This shows that extracardiac pulsations can be easily confused with the anterograde heartbeat. It does not seem realistic to assume that the relatively weak insect heart, and not the 100- to 500-fold more powerful extracardiac system of abdominal pump, could be at all responsible for selective accumulation of haemolymph in anterior parts of the body, for inflation of wings or enhancement of tracheal ventilation.It has been established that thermography from the pericardial region is not specific for the heartbeat. It records subepidermal movement of haemolymph resulting from the actions of both dorsal vessel and extracardiac pressure pulses as well. Shortly before adult eclosion the cardiac and extracardiac pulsations occasionally strike in concert, which profoundly increases the flow of haemolymph through pericardial and perineural sinuses. The relatively strong extracardiac pulsations cause passive movements of various visceral organs, tissue membranes, or tissue folds, giving thus a false impression of an authentic pulsation of tissues. In addition, extracardiac pulsations cause rhythmical movements of haemolymph between various organs, thus preventing haemolymph occlusion at the sites where the heart does not reach. It has been emphasized, finally, that the function of the autonomic nervous system (coelopulse), which integrates extracardiac pulsations, depends on homeostatic moderation of excessive or deficient conditions in insect respiration and haemolymph circulation.     

Why do adult insects not moult?

Hypotheses are presented concerning why mayflies moult after functional wings develop and why most insects cease to moult at this time. The pattern of retention or loss of the subimaginal moult in extant mayflies suggests that this moult may be necessary to complete elongation of caudal filaments and forelegs of adults. It is then analogous to the pupal moult of holometabolous insects. I propose that selection for wing efficiency has normally confined the presence of functional wings to one instar only, which is also the last and reproductive stage. Selection for light wings has generally caused the epidermis of membranous insect wings to degenerate, thereby precluding otherwise advantageous adult moults.     

周期性羽化昆虫形成机制

当昆虫类群表现为长生命周期k(k > 1)年时,成虫的羽化表现为非周期性、周期性和过渡周期性3种形式。非周期性即为成虫每年均羽化,周期性即为成虫每k年才羽化1次,过渡周期性为非周期昆虫逐渐进化为羽化周期性的必经阶段,不同年份羽化的同生群在密度上产生了显著差异,形成了小同生群和优势同生群。自然界中表现出完全羽化周期性的昆虫种类是较少的,但由于其高种群密度的成虫同步性羽化现象,对比非周期性昆虫更易暴发成灾。为明确周期性昆虫演化进程并为林区虫害防控提供理论指导,总结了周期性昆虫的种类和生活特性,不同年间的气候异质性、自然灾害、扩散到未分布区域、天敌、种间和种内竞争等因素均有可能成为过渡周期现象形成的最初驱动力,生活史延长、寄主-天敌互作、低温驱动效应、天敌不敏感-捕食者饱足效应、种间和种内竞争等是促使昆虫羽化周期性形成的可能机制。在林区管理实践中,应提前评估害虫羽化周期性产生的趋势和程度。当成虫表现出完全羽化周期性,应在集中羽化年份内采取见效快的综合防控策略,降低唯一同生群密度至经济阈值以下。当成虫表现出过渡周期性,应加大优势同生群防治力度、降低小同生群防治频率,以及采取天敌林间释放和保育技术以平衡天敌对目标害虫的不同发育阶段种群的控制作用大小,遏制或减缓天敌-寄主互作驱动下的周期性演化进程,逐渐实现由过渡周期阶段向非周期性的逆转。当成虫表现出非周期性,应减少专化性天敌的释放和针对害虫特定阶段的防治措施使用频率,优先选择作用于所有发育阶段且致死率不存在显著差异的防治手段,避免因人为干扰产生的周期性演化和进一步的成灾。     

Use of habitat odour by host‐seeking insects

Locating suitable feeding or oviposition sites is essential for insect survival. Understanding how insects achieve this is crucial, not only for understanding the ecology and evolution of insect–host interactions, but also for the development of sustainable pest‐control strategies that exploit insects' host‐seeking behaviours. Volatile chemical cues are used by foraging insects to locate and recognise potential hosts but in nature these resources usually are patchily distributed, making chance encounters with host odour plumes rare over distances greater than tens of metres. The majority of studies on insect host‐seeking have focussed on short‐range orientation to easily detectable cues and it is only recently that we have begun to understand how insects overcome this challenge. Recent advances show that insects from a wide range of feeding guilds make use of ‘habitat cues’, volatile chemical cues released over a relatively large area that indicate a locale where more specific host cues are most likely to be found. Habitat cues differ from host cues in that they tend to be released in larger quantities, are more easily detectable over longer distances, and may lack specificity, yet provide an effective way for insects to maximise their chances of subsequently encountering specific host cues. This review brings together recent advances in this area, discussing key examples and similarities in strategies used by haematophagous insects, soil‐dwelling insects and insects that forage around plants. We also propose and provide evidence for a new theory that general and non‐host plant volatiles can be used by foraging herbivores to locate patches of vegetation at a distance in the absence of more specific host cues, explaining some of the many discrepancies between laboratory and field trials that attempt to make use of plant‐derived repellents for controlling insect pests.     

Advances in Transgenic Research for Insect Resistance in Sugarcane

The first phase of transgenic research in sugarcane concentrated on the development and evaluation of transgenic lines transformed for resistance to biotic stresses, particularly diseases and insect pests. Sugarcane is attacked by a range of insects including tissue borers, sucking pests and canegrubs. Losses due to these pests are estimated to be around 10%. Although chemical control and integrated pest management are regularly practiced for the control of insect pests, success is often limited due to practical difficulties. The genetic complexity of sugarcane coupled with the non-availability of resistance genes in the germplasm has made conventional breeding for insect resistance difficult. In this context, transgenic technology has become a handy tool for imparting insect resistance to an elite variety which is otherwise superior for most other agronomic traits. A number of transgenic sugarcane lines have been developed with genes expressing Cry proteins, proteinase inhibitors or lectins resistant to borers, sucking insects or grubs. While commercializing transgenic lines, issues such as higher and stable transgene expression, preparedness for resistance management and non-target effects need to be addressed. To manage the constant threat of resistance development in target insects, it is imperative to deploy field-level strategies taking clues from other crops coupled with the search for new potent replacement molecules for transformation.     

杀虫剂抗性: 遗传学、基因组学及应用启示

杀虫剂抗性已成为害虫防治工作需要解决的一个重要问题,也是一种人为的、自然选择的重要的进化现象,开展抗药性的研究不仅为抗性的监测、治理和农药工业的发展提供科学参考,还可以揭示生物进化的一些基本规律。在过去的10年,昆虫对许多化学杀虫剂抗药性的分子基础得到了进一步阐明,已从果蝇Drosophila melanogaster中克隆了杀虫剂的靶标基因,还查明了一些害虫的与抗性相关联的基因突变。最近,随着经注释的昆虫基因组的出现,由复杂多基因酶系如酯酶、细胞色素P450酶及谷胱甘肽S-转移酶介导的抗性的机制有了突破性的进展,有关杀虫剂抗性的进化以及抗性基因的传播模式也逐步得到揭示。基因组技术在揭示昆虫其他可能的抗药性机制以及在发现新的杀虫剂靶标方面将发挥更大的作用。     

Mechanics and aerodynamics of insect flight control

Insects have evolved sophisticated fight control mechanisms permitting a remarkable range of manoeuvres. Here, I present a qualitative analysis of insect flight control from the perspective of flight mechanics, drawing upon both the neurophysiology and biomechanics literatures. The current literature does not permit a formal, quantitative analysis of flight control, because the aerodynamic force systems that biologists have measured have rarely been complete and the position of the centre of gravity has only been recorded in a few studies. Treating the two best-known insect orders (Diptera and Orthoptera) separately from other insects, I discuss the control mechanisms of different insects in detail. Recent experimental studies suggest that the helicopter model of flight control proposed for Drosophila spp. may be better thought of as a facultative strategy for flight control, rather than the fixed (albeit selected) constraint that it is usually interpreted to be. On the other hand, the so-called 'constant-lift reaction' of locusts appears not to be a reflex for maintaining constant lift at varying angles of attack, as is usually assumed, but rather a mechanism to restore the insect to pitch equilibrium following a disturbance. Differences in the kinematic control mechanisms used by the various insect orders are related to differences in the arrangement of the wings, the construction of the flight motor and the unsteady mechanisms of lift production that are used. Since the evolution of insect flight control is likely to have paralleled the evolutionary refinement of these unsteady aerodynamic mechanisms, taxonomic differences in the kinematics of control could provide an assay of the relative importance of different unsteady mechanisms. Although the control kinematics vary widely between orders, the number of degrees of freedom that different insects can control will always be limited by the number of independent control inputs that they use. Control of the moments about all three axes (as used by most conventional aircraft) has only been proven for larger flies and dragonflies, but is likely to be widespread in insects given the number of independent control inputs available to them. Unlike in conventional aircraft, however, insects' control inputs are likely to be highly non-orthogonal, and this will tend to complicate the neural processing required to separate the various motions.     

Host plant resistance for insect control in some important crop plants

Insect pests of cereal crop plants are among the most destructive and devastating factors limiting world food production. Insects often inflict losses of 15 to 50% of the yield of some crops in various parts of the world. Insects also provide infection courts for various pathogens that inflict even greater damage. The nature of these losses is especially tragic for poor farmers in the tropics since the insects consume dry matter already produced using limited resources. Reduction of these losses would enhance the world food supply available to man, even if actual production remains the same, by allotting to man and animals that proportion of production now consumed by insects. Control of insect‐inflicted damage on a world‐wide basis is a very difficult task. Chemical control is effective and readily available in some, but not all, countries. However, chemical control is often unavailable, ineffective, too expensive, and likely to create environmental and safety hazards that make it unsuitable for many parts of the world. Several excellent examples of biological control of insects also attest to the potential usefulness of this method of reducing insect‐inflicted losses, but this method also has been ineffective for certain major insect pests due to lack of effective identification, distribution, and maintenance of biological control agents. On the whole, host plant resistance (HPR) to insects has been the most successful and widely used method of control. HPR has been used successfully in a number of major crop species to help control damage inflicted by major insect pests. However, problems with identifying sources of resistance and transferring it to usable varieties have often been difficult to overcome. Also, once resistance is deployed, changes in insect populations which allow them to overcome the resistance sometimes eliminate the potential advantages of resistant varieties. However, methods to quantify host‐insect interactions have improved significantly over the past few years, and the prospects for developing and maintaining usable levels of resistance to several major insect pests of most major crops are good. Studies of the biochemical and biophysical mechanisms of insect resistance have added a new dimension to HPR work. Knowledge of specific factors that contribute to insect resistance can be extremely useful to plant breeders and entomologists. The ability to identify and select for one specific plant component or group of components can speed the development of resistant varieties. Studies of the effect of such resistance components on insect development and behavior can help determine the likelihood of insects being able to overcome the resistance factors. Studies of the effect of the resistance factors on host plant yield, performance, or quality may help plant breeders overcome the problems often associated with the development of high yielding and high quality resistant varieties. Identification of several different mechanisms of resistance could provide breeders the opportunity to adjust levels of several resistance components in order to arrive at a proper blend of resistance, quality, and yield. Furthermore, development of varieties with multiple mechanisms of resistance should slow the development of new biotypes of insects that could overcome the HPR since the insects would have to simultaneously or sequentially develop the ability to overcome several resistance factors. Therefore, knowledge of mechanisms of insect resistance could facilitate the development of more stable and long‐lasting types of resistance.     

昆虫的RNA干扰

RNA干扰(RNAi)是一种强有力的分子生物学技术, 在昆虫研究中得到了较多的应用。目前, RNAi技术主要应用于昆虫功能基因和功能基因组研究, 已在多个目的19种昆虫上实现了RNAi。在昆虫上实现RNAi的方法主要有注射、浸泡、喂食、转基因和病毒介导等方法, 这些方法各有特点, 其中喂食法因其简单而最有应用前景。昆虫RNAi的系统性较为复杂, 只有部分昆虫具有RNAi的系统性。昆虫中RNAi信号传导的基因可能是sid-1, 但昆虫RNAi的系统性机理还不是很清楚。转基因植物产生的dsRNA实现了对作物的保护, 证实了RNAi技术可用于害虫控制, 为害虫控制开辟了新领域。昆虫的RNAi研究处在起步阶段, 研究昆虫RNAi的机理, 特别是RNAi在昆虫体内的系统性扩散机理, 改进实现RNAi的方法, 提高RNAi技术在昆虫研究中的应用, 有利于昆虫基因功能鉴定和害虫控制, 促进昆虫学科的发展。     

Feasibility,limitation and possible solutions of RNAi‐based technology for insect pest control

Abstract Numerous studies indicate that target gene silencing by RNA interference (RNAi) could lead to insect death. This phenomenon has been considered as a potential strategy for insect pest control, and it is termed RNAi‐mediated crop protection. However, there are many limitations using RNAi‐based technology for pest control, with the effectiveness target gene selection and reliable double‐strand RNA (dsRNA) delivery being two of the major challenges. With respect to target gene selection, at present, the use of homologous genes and genome‐scale high‐throughput screening are the main strategies adopted by researchers. Once the target gene is identified, dsRNA can be delivered by micro‐injection or by feeding as a dietary component. However, micro‐injection, which is the most common method, can only be used in laboratory experiments. Expression of dsRNAs directed against insect genes in transgenic plants and spraying dsRNA reagents have been shown to induce RNAi effects on target insects. Hence, RNAi‐mediated crop protection has been considered as a potential new‐generation technology for pest control, or as a complementary method of existing pest control strategies; however, further development to improve the efficacy of protection and range of species affected is necessary. In this review, we have summarized current research on RNAi‐based technology for pest insect management. Current progress has proven that RNAi technology has the potential to be a tool for designing a new generation of insect control measures. To accelerate its practical application in crop protection, further study on dsRNA uptake mechanisms based on the knowledge of insect physiology and biochemistry is needed.     

Insect quality control: synchronized sex,mating system,and biological rhythm

The sterile insect technique (SIT) is a method of eradicating insects by releasing mass-reared sterilized males into fields to reduce the hatchability of eggs laid by wild females that have mated with the sterile males. SIT requires mass-production of the target insect, and maintenance of the quality of the mass-reared insects. The most important factor is successful mating between wild females and sterile males because SIT depends on their synchronized copulation. Therefore, understanding the mating systems and fertilization processes of target insects is prerequisite. Insect behavior often has circadian rhythms that are controlled by a biological clock. However, very few studies of relationships between sterile insect quality and circadian rhythm have been performed compared with the amount of research on the mating ability of target insects. The timing of male copulation attempts with receptivity of females is key to successful mating between released males and wild females. Therefore, we should focus on the mechanisms controlling the timing of mating in target insects. On the other hand, in biological control projects, precise timing of the release of natural enemies to attack pest species is required because behavior of pests and control agents are affected by their circadian rhythms. Involving both chronobiologists and applied entomologists might produce novel ideas for sterile insect quality control by synchronized sex between mass-reared and wild flies, and for biological control agent quality by matching timing in activity between predator activity and prey behavior. Control of the biological clocks in sterile insects or biological control agents is required for advanced quality control of rearing insects.     

Photic entrainment of circadian rhythms by illumination of implanted brain tissues in the cockroach Blaberus craniifer

The locomotor activity rhythms of ‘blinded’ Blaberus craniifer adults receiving transplants of either an entire brain or both optic lobes of a donor B. craniifer were studied in contrast to the activity rhythms of surgical control and untreated insects. When the implants were located in the abdomen of a ‘host’ insect beneath a small glass ‘window’ and exposed to a 12L : 12D photoperiod entrainment occurred and the activity of such ‘host’ insects receiving either type of implant was comparable to that of control insects. The results provide some substantiating evidence that locomotor activity rhythms may be controlled by a hormonal clock.     

High-nickel insects and nickel hyperaccumulator plants: A review

Insects can vary greatly in whole‐body elemental concentrations. Recent investigations of insects associated with Ni hyperaccumulator plants have identified insects with relatively elevated whole‐body Ni levels. Evaluation of the limited data available indicates that a whole‐body Ni concentration of 500 μg Ni/g is exceptional: I propose that an insect species with a mean value of 500 μg Ni/g or greater, in either larval/nymphal or adult stages, be considered a “high‐Ni insect”. Using the 500 μg Ni/g criterion, 15 species of high‐Ni insects have been identified to date from studies in Mpumalanga (South Africa), New Caledonia and California (USA). The highest mean Ni concentration reported is 3 500 μg Ni/g for nymphs of a South African Stenoscepa species (Orthoptera: Pyrgomorphidae). The majority of high‐Ni insects (66%) are heteropteran herbivores. Studies of high‐Ni insect host preference indicate they are monophagous (or nearly so) on a particular Ni hyperaccumulator plant species. Much of the Ni in bodies of these insects is in their guts (up to 66%–75%), but elevated levels have also been found in Malpighian tubules, suggesting efficient elimination as one strategy for dealing with a high‐Ni diet. Tissue levels of Ni are generally much lower than gut concentrations, but up to 1200 μg Ni/g has been reported from exuviae, suggesting that molting may be another pathway of Ni elimination. One ecological function of the high Ni concentration of these insects may be to defend them against natural enemies, but to date only one experimental test has supported this “elemental defense” hypothesis. Community‐level studies indicate that high‐Ni insects mobilize Ni into food webs but that bioaccumulation of Ni does not occur at either plant‐herbivore or herbivore‐predator steps. Unsurprisingly, Ni bioaccumulation indices are greater for high‐Ni insects compared to other insect species that feed on Ni hyperaccumulator plants. There is some evidence of Ni mobilization into food webs by insect visitors to flowers of Ni hyperaccumulator plants, but no high‐Ni insect floral visitors have been reported.     

Sexually transmitted diseases of insects: distribution, evolution, ecology and host behaviour

Sexually transmitted diseases (STDs) of insects are known from the mites, nematodes, fungi, protists and viruses. In total 73 species of parasite and pathogen from approximately 182 species of host have been reported. Whereas nearly all vertebrate STDs are viruses or bacteria, the majority of insect STDs are multicellular ectoparasites, protistans or fungi. Insect STDs display a range of transmission modes, with 'pure' sexual transmission only described from ectoparasites, all of which are mites, fungi or nematodes, whereas the microparasitic endo-parasites tend to show vertical as well as sexual transmission. The distribution of STDs within taxa of insect hosts appears to be related to the life histories of the hosts. In particular, STDs will not be able to persist if host adult generations do not overlap unless they are also transmitted by some alternative route. This explains the observation that the Coleoptera seem to suffer from more STDs than other insect orders, since they tend to diapause as adults and are therefore more likely to have overlapping generations of adults in temperate regions. STDs of insects are often highly pathogenic, and are frequently responsible for sterilizing their hosts, a feature which is also found in mammalian STDs. This, combined with high prevalences indicates that STDs can be important in the evolution and ecology of their hosts. Although attempts to demonstrate mate choice for uninfected partners have so far failed it is likely that STDs have other effects on host mating behaviour, and there is evidence from a few systems that they might manipulate their hosts to cause them to mate more frequently. STDs may also play a part in sexual conflict, with males in some systems possibly gaining a selective advantage from transmitting certain STDs to females. STDs may well be important factors in host population dynamics, and some have the potential to be useful biological control agents, but empirical studies on these subjects are lacking.     

昆虫钠离子通道的研究进展

昆虫只有一个或两个电压门控钠离子通道α亚基基因,但两种转录后修饰(选择性剪切和RNA编辑)实现了昆虫钠离子通道的功能多样性.昆虫β辅助亚基TipE和TEH1-4在钠离子通道表达和调控中也起着重要作用.电压门控钠离子通道在动作电位的产生和传递中至关重要,是多种天然和人工合成神经毒素及杀虫剂的作用靶标,包括广泛使用的拟除虫...     

Rapid induction of immune density-dependent prophylaxis in adult social insects

The innate immune system provides defence against parasites and pathogens. This defence comes at a cost, suggesting that immune function should exhibit plasticity in response to variation in environmental threats. Density-dependent prophylaxis (DDP) has been demonstrated mostly in phase-polyphenic insects, where larval group size determines levels of immune function in either adults or later larval instars. Social insects exhibit extreme sociality, but DDP has been suggested to be absent from these ecologically dominant taxa. Here we show that adult bumble-bee workers (Bombus terrestris) exhibit rapid plasticity in their immune function in response to social context. These results suggest that DDP does not depend upon larval conditions, and is likely to be a widespread and labile response to rapidly changing conditions in adult insect populations. This has obvious ramifications for experimental analysis of immune function in insects, and serious implications for our understanding of the epidemiology and impact of pathogens and parasites in spatially structured adult insect populations.     

Analysis of nubbin expression patterns in insects

Previous studies have shown that the gene nubbin (nub) exhibits large differences in expression patterns between major groups of arthropods. This led us to hypothesize that nub may have evolved roles that are unique to particular arthropod lineages. However, in insects, nub has been studied only in Drosophila. To further explore its role in insects in general, we analyzed nub expression patterns in three hemimetabolous insect groups: zygentomans (Thermobia domestica, firebrat), dyctiopterans (Periplaneta americana, cockroach), and hemipterans (Oncopeltus fasciatus, milkweed bug). We discovered three major findings. First, observed nub patterns in the ventral central nervous system ectoderm represent a synapomorphy (shared derived feature) that is not present in other arthropods. Furthermore, each of the analyzed insects exhibits a species-specific nub expression in the central nervous system. Second, recruitment of nub for a role in leg segmentation occurred early during insect evolution. Subsequently, in some insect lineages (cockroaches and flies), this original role was expanded to include joints between all the leg segments. Third, the nub expression in the head region shows a coordinated change in association with particular mouthpart morphology. This suggests that nub has also gained an important role in the morphological diversification of insect mouthparts. Overall, the obtained data reveal an extraordinary dynamic and diverse pattern of nub evolution that has not been observed previously for other developmental genes.     

Mutations and their use in insect control

Traditional chemically based methods for insect control have been shown to have serious limitations, and many alternative approaches have been developed and evaluated, including those based on the use of different types of mutation. The mutagenic action of ionizing radiation was well known in the field of genetics long before it was realized by entomologists that it might be used to induce dominant lethal mutations in insects, which, when released, could sterilize wild female insects. The use of radiation to induce dominant lethal mutations in the sterile insect technique (SIT) is now a major component of many large and successful programs for pest suppression and eradication. Adult insects, and their different developmental stages, differ in their sensitivity to the induction of dominant lethal mutations, and care has to be taken to identify the appropriate dose of radiation that produces the required level of sterility without impairing the overall fitness of the released insect. Sterility can also be introduced into populations through genetic mechanisms, including translocations, hybrid incompatibility, and inherited sterility in Lepidoptera. The latter phenomenon is due to the fact that this group of insects has holokinetic chromosomes. Specific types of mutations can also be used to make improvements to the SIT, especially for the development of strains for the production of only male insects for sterilization and release. These strains utilize male translocations and a variety of selectable mutations, either conditional or visible, so that at some stage of development, the males can be separated from the females. In one major insect pest, Ceratitis capitata, these strains are used routinely in large operational programs. This review summarizes these developments, including the possible future use of transgenic technology in pest control.