蜕皮激素和IIS-TORC1信号的分子互作

蜕皮激素信号主导调控昆虫的蜕皮和变态,决定昆虫的发育时间;IIS-TORC1信号整合生长因子、激素、营养和能量信号,决定昆虫的生长速率。蜕皮激素和IIS-TORC1信号之间发生3种分子互作:(1)IIS-TORC1信号促进前胸腺和卵巢合成蜕皮激素前体。(2)在蜕皮和变态期间,蜕皮激素抑制脂肪体细胞内IIS-TORC1信号、Myc的转录、细胞生长及其内分泌功能,导致脑神经分泌细胞分泌胰岛素样肽的功能减弱,从而降低昆虫全身性的IIS-TORC1信号。(3)在幼虫摄食期间,胰岛素信号抑制FOXO的转录活性,降低了蜕皮激素受体EcR的转录共激活因子DOR编码基因的转录水平,从而阻碍了蜕皮激素信号传导。蜕皮激素信号和IIS-TORC1信号协同调控发育时间和生长速率共同决定昆虫的个体大小。     

昆虫变态发育的激素和营养调控研究进展与展望

变态发育是促进昆虫进化和多样性形成的重要因素之一。昆虫变态发育主要受到蜕皮激素和保幼激素的协同调控,通过信号通路诱导下游基因表达,保证蜕皮、组织重塑等生理过程的正确发生。除内在激素外,外在营养物质亦可通过营养信号影响激素信号进而控制变态发育进程。本文主要综述了近十年来我国科研工作者在昆虫变态发育的激素和营养调控机制研究方面所取得的突出成果,并对未来潜在的研究方向进行了展望,以期对我国的害虫防治和益虫利用研究提供理论指导。     

昆虫脂肪体的代谢作用

<正> 昆虫体内蕴藏着大量的脂肪体。这些脂肪体从表面上看似乎只是类脂物的贮存,但实际上是昆虫生长、发育、变态和生殖等代谢活动的中心组织。由于脂肪体能贮存营养、解毒及为昆虫生命的周期活动提供各种生物合成的代谢产物,因此,人们将昆虫的脂肪体比拟为脊椎动物的肝脏。昆虫脂肪体的代谢作用是受激素调节控制的,本文介绍这方面研究结果的概况。 1.脂肪体的结构 昆虫的脂肪体有不同的形态,不同目的昆虫脂肪体的结构不同:有纸片状(半翅目)、绳子状(直翅目)、带状(鳞翅目)和球状(鞘翅目)     

昆虫蜕皮激素信号转导途径研究进展

蜕皮与变态是全变态昆虫典型的发育特征。调控昆虫蜕皮与变态的激素主要有蜕皮激素和保幼激素。目前已经阐明了蜕皮激素的核受体EcR及部分核信号转导途径,但蜕皮激素是否存在膜受体及膜信号转导途径研究很少。研究证明,蜕皮激素存在细胞质中的信号转导分子和途径,蜕皮激素通过NTF2和Ran调控EcR入核启动基因转录。蜕皮激素使细胞质中的热休克蛋白Hsc70部分入核与USP结合启动基因转录。蜕皮激素通过蛋白激酶PKC使伴侣蛋白calponin磷酸化,参与蜕皮激素信号途径的基因转录。这些研究结果说明蜕皮激素除了有核受体和核受体信号转导途径外,还存在细胞膜受体和细胞膜信号转导途径。     

昆虫的变态发育研究

昆虫变态发育使得昆虫成为地球陆地上种类最多、数量最大、分布最广、生活环境最多样化的一群生物。变态使昆虫在其生命周期中的不同发育时期表现出完全不同的形态、结构、功能和生活习性的变化,有利于昆虫迁飞转移,扩大其求偶交配、生活和生存环境空间。昆虫变态发育的变化是长期自然环境适应、协同进化的结果,受激素、营养和基因的精确调控。本文简要介绍了昆虫变态的类型、激素调控、营养调控和基因调控方面的研究进展,以及研究昆虫变态发育的科学和应用意义。     

昆虫激素作用原理概述

经过近40年来的不断实验和研究,已证实昆虫体内具有控制个体生长、发育、蜕皮、变态以及生殖等生理过程的内分泌系统,内分泌器官分泌出来的激素(内激素),借体液运送,作用于体内某些器宫,引起相应的生理变化过程。本文首先扼要地介绍一下关于脑激素、蜕皮激素和保幼激素在调节控制昆虫胚后发育过程中的生理作用,然后进一步概述蜕皮激素和保幼激素是如何起作用的。     

昆虫胰岛素生理功能的研究进展

胰岛素是由胰岛β细胞经过一些内源性或外源性物质的诱发而分泌的蛋白质激素,通过细胞的信号转导控制调节糖、脂肪和蛋白质代谢,从而影响生殖以及衰老等生长发育过程。近年来研究陆续证实该途径在昆虫中也普遍存在。本文综述了昆虫胰岛素信号途径,胰岛素在调控昆虫生长发育、生殖和行为方面的重要作用,并且胰岛素可以通过对保幼激素和蜕皮激素的影响间接作用于昆虫,对了解胰岛素在昆虫生理功能中的作用及害虫综合防治策略的制定具有积极意义。     

昆虫变态发育过程中的细胞自噬和凋亡

在昆虫变态期,幼虫组织发生退化或消亡,原因在于蜕皮甾醇激素（ecdysteroid）,即通常所说的蜕皮激素,诱导这些组织的细胞发生了自噬(autophagy)和凋亡(apoptosis)的程序性细胞死亡(programmed cell death,PCD)。一般情况下,自噬途径构成一种饥饿应激适应性以避免细胞的死亡,表现为低水平Cvt泡(Cvt vesicle)和自噬体(autophagosome)对部分胞质溶胶、蛋白聚集体和细胞器的吞噬和降解。昆虫进入变态发育时,由于蜕皮激素的激活,由遗传级联系统调控的PCD机制被启动,低水平的常态自噬转入高水平的自噬并同时诱发凋亡,细胞进入不可逆的死亡,导致幼虫组织在变态期退化或消亡。对果蝇Drosophila变态期PCD机制中最重要的发现是:（1）在自噬发生的PI3KⅠ- Tor 和 PI3KⅢ的分子通路中,由自噬相关蛋白Atg1引发的高水平自噬能够诱导凋亡;（2）蜕皮激素诱导表达的βFTZ-F1,E93,BR-C,E74A等转录因子不但激活凋亡的Caspases通路,还能诱导自噬的发生。     

蜜蜂脂肪体的形态和功能研究进展

脂肪体是昆虫体内的一种多功能器官,近似于脊椎动物的肝脏, 分布于昆虫腹部、胸部甚至头部腔体中,以腹部脂肪体最为发达。蜜蜂脂肪体有外周脂肪体和围脏脂肪体两种类型,由营养细胞、尿酸盐细胞和绛色细胞组成。同其他昆虫中类似,脂肪体在蜜蜂的生命活动中扮演着重要的角色,其形态和功能随发育阶段、季节和劳动分工的变化而变化。脂肪体结构相对简单,但生理功能非常复杂。脂肪体最主要的功能是能量物质的储存和代谢,其不仅是蜜蜂营养物质(即脂质、碳水化合物和蛋白质)的中央储存库,而且是营养代谢的中间站,具有多种能量和物质相互转换的酶系,承担代谢水的供应并合成嘌呤和嘧啶及许多重要的蛋白质。同时,脂肪体是昆虫发育和行为调控过程中各种激素和营养信号的交换中心,脂肪体激素和营养信号参与调控蜜蜂脂肪体发育、营养物质代谢、生殖及劳动分工。脂肪体兼具能量储存和释放、生物合成和分解、营养感知调节、代谢信号整合、内分泌调节、免疫和解毒、磁场感受、提高抗寒能力、保护体腔内器官等多种功能。鉴于脂肪体的重要作用,蜜蜂脂肪体形态和功能的研究成果可以对昆虫营养信号通路的解析、蜂产品高产良种的选育和蜜蜂病害防治的研究提供参考和思路。     

昆虫变态发育类型与调控机制

昆虫变态发育使得昆虫成为地球上种类最多、分布最广的动物种群。它特指末龄幼虫化蛹,或者蛹向成虫的转变过程。根据变态剧烈程度,可将昆虫变态发育简单分为增节变态、表变态、原变态、不完全变态及全变态5种类型。此外,昆虫变态发育是一个极其复杂的生物学过程,受到激素、基因、营养等多种因素的精密调控。本文简要介绍了昆虫变态发育的类型和分子调控机理方面的研究进展。     

Hormonal and nutritional regulation of insect fat body development and function

The insect fat body is an organ analogue to vertebrate adipose tissue and liver and functions as a major organ for nutrient storage and energy metabolism. Similar to other larval organs, fat body undergoes a developmental “remodeling” process during the period of insect metamorphosis, with the massive destruction of obsolete larval tissues by programmed cell death and the simultaneous growth and differentiation of adult tissues from small clusters of progenitor cells. Genetic ablation of Drosophila fat body cells during larval‐pupal transition results in lethality at the late pupal stage and changes sizes of other larval organs indicating that fat body is the center for pupal development and adult formation. Fat body development and function are largely regulated by several hormonal (i.e. insulin and ecdysteroids) and nutritional signals, including oncogenes and tumor suppressors in these pathways. Combining silkworm physiology with fruitfly genetics might provide a valuable system to understand the mystery of hormonal regulation of insect fat body development and function. © 2009 Wiley Periodicals, Inc.     

Prothoracicotropic hormone regulates developmental timing and body size in Drosophila

In insects, control of body size is intimately linked to nutritional quality as well as environmental and genetic cues that regulate the timing of developmental transitions. Prothoracicotropic hormone (PTTH) has been proposed to play an essential role in regulating the production and/or release of ecdysone, a steroid hormone that stimulates molting and metamorphosis. In this report, we examine the consequences on Drosophila development of ablating the PTTH-producing neurons. Surprisingly, PTTH production is not essential for molting or metamorphosis. Instead, loss of PTTH results in delayed larval development and eclosion of larger flies with more cells. Prolonged feeding, without changing the rate of growth, causes the overgrowth and is a consequence of low ecdysteroid titers. These results indicate that final body size in insects is determined by a balance between growth-rate regulators such as insulin and developmental timing cues such as PTTH that set the duration of the feeding interval.     

Towards a mechanistic understanding of insect life history evolution: oxygen‐dependent induction of moulting explains moulting sizes

Moults characterise insect growth trajectories, typically following a consistent pattern known as Dyar's rule; proportional size increments remain constant across inter‐instar moults. Empirical work suggests that oxygen limitation triggers moulting. The insect respiratory system, and its oxygen supply capacity, grows primarily at moults. It is hypothesized that the oxygen demand increases with increasing body mass, eventually meeting the oxygen supply capacity at an instar‐specific critical mass where moulting is triggered. Deriving from this hypothesis, we develop a novel mathematical model for moulting and growth in insect larvae. Our mechanistic model has great success in predicting moulting sizes in four butterfly species, indirectly supporting a size‐dependent mechanism underlying moulting. The results demonstrate that an oxygen‐dependent induction of moulting mechanism would be sufficient to explain moulting sizes in the study species. Model predictions deviated slightly from Dyar's rule, the deviations being typically negligible within the present data range. The developmental decisions (e.g. moulting) made by growing larvae significantly affect age and size at maturity, which has important life history implications. The successful modelling of moulting presented here provides a novel framework for the development of realistic insect growth models, which are required for a better understanding of life history evolution.     

The physiological basis of reaction norms: the interaction among growth rate, the duration of growth and body size

The general effects of temperature and nutritional quality ongrowth rate and body size are well known. We know little, however,about the physiological mechanisms by which an organism translatesvariation in diet and temperature into reaction norms of bodysize or development time. We outline an endocrine-based physiologicalmechanism that helps explain how this translation occurs inthe holometabolous insect Manduca sexta (Sphingidae). Body sizeand development time are controlled by three factors: (i) growthrate, (ii) the timing of the cessation of juvenile hormone secretion(measured by the critical weight) and (iii) the timing of ecdysteroidsecretion leading to pupation (the interval to cessation ofgrowth [ICG] after reaching the critical weight). Thermal reactionnorms of body size and development time are a function of howthese three factors interact with temperature. Body size issmaller at higher temperatures, because the higher growth ratedecreases the ICG, thereby reducing the amount of mass thatcan accumulate. Development time is shorter at higher temperaturesbecause the higher growth rate decreases the time required toattain the critical weight and, independently, controls theduration of the ICG. Life history evolution along altitudinal,latitudinal and seasonal gradients may occur through differentialselection on growth rate and the duration of the two independentlycontrolled determinants of the growth period.     

Chronic malnutrition favours smaller critical size for metamorphosis initiation in Drosophila melanogaster

Critical size at which metamorphosis is initiated represents an important checkpoint in insect development. Here, we use experimental evolution in Drosophila melanogaster to test the long-standing hypothesis that larval malnutrition should favour a smaller critical size. We report that six fly populations subject to 112 generations of laboratory natural selection on an extremely poor larval food evolved an 18% smaller critical size (compared to six unselected control populations). Thus, even though critical size is not plastic with respect to nutrition, smaller critical size can evolve as an adaptation to nutritional stress. We also demonstrate that this reduction in critical size (rather than differences in growth rate) mediates a trade-off in body weight that the selected populations experience on standard food, on which they show a 15-17% smaller adult body weight. This illustrates how developmental mechanisms that control life history may shape constraints and trade-offs in life history evolution.     

Gamete production and sexual size dimorphism in an insect (Orchesella cincta) with indeterminate growth

Abstract 1. The relationship of growth and body size with reproductive effort in animal species has been studied much less for males than for females. This imbalance applies to Orchesella cincta (L.) (Collembola), an insect with indeterminate growth, in which egg production is related positively to body size and negatively to growth.
2. To allow a comparison of the reproductive effort of male and female O. cincta , development and growth in immature stages of both sexes, and growth and spermatophore production for adult males were studied.
3. Embryonic development time and hatchling size did not differ between prospective males and females, but from hatching on the trajectories diverged, with males growing more slowly and maturing earlier and at a much smaller body size than females.
4. Neither the number of spermatophores deposited in the first adult instar (= inter-moult period) nor the total number of spermatophores deposited during seven instars was related to body size or growth.
5. Differences in growth rate between instars with and without spermatophore deposition indicated that the physiology of spermatophore production inhibits growth, which, however, was compensated for during the next instar.
6. The difference in the relationship of gamete production with body size and growth between males and females explains the divergence of their size at maturity.     

Body size determination in insects: a review and synthesis of size‐ and brain‐dependent and independent mechanisms

Body size determination requires a mechanism for sensing size and a mechanism for linking size information to the termination of growth. Although the hormonal mechanisms that terminate growth are well elucidated, the mechanisms by which a body senses its own size are only partially understood; most of this understanding has come from the study of the mechanisms that control insect moulting and metamorphosis. We first review and discuss advances in our understanding of the physiological mechanisms by which insect larvae sense their size. Second, we present new findings on how larvae in which the size‐sensing mechanism has been disrupted eventually terminate growth (in a size‐independent manner). We synthesize recent insights into the genetic and molecular mechanisms of ecdysteroid regulation in Drosophila melanogaster with developmental physiology findings in Manduca sexta, paving the way for an integrated understanding of the mechanisms of body size regulation.     

拟寄生昆虫中的过寄生现象

对拟寄生昆虫同种过寄生现象做了综述。拟寄生昆虫的认识能力和寄生经历、其与寄主数量的相对比例、寄主的大小及两次被寄生的时间间隔等是导致过寄生是否出现的主要因素。过寄生常导致拟寄生昆虫发育历期延长、存活率下降、个体变小、子代雌性比降低。试验研究和大量饲养中应采取措施避免过寄生。     

Tradeoffs between somatic and gonadal investments during development in the African clawed frog (Xenopus laevis)

Tradeoffs between time to and size at metamorphosis occur in many organisms with complex life histories. The ability to accelerate metamorphosis can increase survival to the next life stage, but the resulting smaller size at metamorphosis is often associated with lower post-metamorphic survival or reduced fecundity of adults. Reduced fecundity is thought to be because of reduced energy reserves, longer time to maturity, or reduced capacity to carry eggs or compete for mates. This pattern could also be explained by a shift in allocation to somatic growth that further retards the growth or development of reproductive tissues. The main goal of this study was to determine if the relationship between growth and development of somatic and gonadal tissues depends on environmental conditions. We address this question through two experiments in which we quantify the development and growth of the body and gonads of Xenopus laevis reared in different resource environments. First, tadpoles were reared communally and development and growth were evaluated over time. Restricted food reduced somatic and gonadal growth rate, but did not affect the developmental rate of either tissue type. Second, tadpoles were reared individually and evaluated at metamorphosis. Restricted food reduced somatic development and growth, but only influenced size, and not developmental stage of testes at metamorphosis. This work demonstrates that environmental conditions influence tradeoffs between growth and development of somatic and gonadal tissues, apparently in a sex-specific manner. These tradeoffs may contribute to phenotypic correlations between small size and reduced fitness.