本文選題：Bergmanns’rule　切入點(diǎn)：果蠅　出處：《浙江大學(xué)》2016年博士論文　論文類型：學(xué)位論文

【摘要】：環(huán)境因素對(duì)個(gè)體發(fā)育的影響是一個(gè)經(jīng)久不衰的研究命題,在眾多因素中,氣候的變更,溫度的變化無處不在,它對(duì)動(dòng)物個(gè)體發(fā)育乃至種群進(jìn)化產(chǎn)生巨大的影響。19世紀(jì),德國(guó)生物學(xué)家Carl Bergmann發(fā)現(xiàn),動(dòng)物個(gè)體的體型與生存地區(qū)緯度呈正相關(guān)關(guān)系,緯度越高,溫度越寒冷的地區(qū),動(dòng)物個(gè)體體型越大,緯度越低,溫度越溫暖的地區(qū),動(dòng)物個(gè)體體型越小。這就是著名的Bergmann's rule,它總結(jié)了生存溫度對(duì)溫血?jiǎng)游矬w型影響的一般規(guī)律。在非脊椎動(dòng)物中,動(dòng)物體形與溫度呈負(fù)相關(guān)的關(guān)系更加普遍存在,我們稱之為溫度體形規(guī)律(Temperature-Size Rule,TSR)。從19世紀(jì)初開始,人們對(duì)各種動(dòng)物進(jìn)行研究和考察,嘗試解釋溫度影響動(dòng)物體型的原因,至今我們對(duì)其分子細(xì)胞機(jī)制仍然知之甚少。模式動(dòng)物果蠅是用來研究溫度影響體型規(guī)律的良好載體,早在1940年,人們已經(jīng)開始利用果蠅研究溫度影響體型的規(guī)律,果蠅在其適宜的生存溫度范圍內(nèi),成熟體的體型大小具有強(qiáng)烈的溫度可塑性,比如低溫環(huán)境(18℃)下飼養(yǎng)的果蠅的體型比在高溫環(huán)境(25℃)下飼養(yǎng)的果蠅大10%左右。果蠅幼蟲階段是發(fā)育積累并決定成熟體體型大小的決定性階段,于是我們主要研究幼蟲的發(fā)育機(jī)制。果蠅幼蟲與成蟲一樣,具有豐富的感知系統(tǒng),包括化學(xué)感知、機(jī)械感知、溫度感知等。在幼蟲外周分布著豐富的初級(jí)感知神經(jīng)元,它們投射到腦中央神經(jīng)元形成簡(jiǎn)單且功能齊全的神經(jīng)網(wǎng)絡(luò)。初級(jí)感知神經(jīng)元是動(dòng)物感知外界環(huán)境因素(溫度)的必經(jīng)之路,我們以研究果蠅幼蟲的溫度感知神經(jīng)系統(tǒng)為立足點(diǎn),以人工操控神經(jīng)元的興奮性為手段,結(jié)合基本的細(xì)胞分子檢測(cè)方法研究溫度影響個(gè)體發(fā)育的機(jī)制。果蠅胰島素是調(diào)節(jié)幼蟲發(fā)育的關(guān)鍵分子之一并且具有物種保守性,主要的幾個(gè)果蠅胰島素由胰島素生成細(xì)胞IPCs (insulin producing cells),一個(gè)分泌性神經(jīng)元,合成與分泌。我們發(fā)現(xiàn)阻斷IPCs可以消除低溫促進(jìn)幼蟲體型增大的影響,而激活I(lǐng)PCs則可以促進(jìn)幼蟲體型增加,通過在體鈣成像實(shí)驗(yàn),我們發(fā)現(xiàn)低溫刺激可以興奮IPCs,通過熒光定量PCR,免疫組織化學(xué)和免疫印記實(shí)驗(yàn),我們發(fā)現(xiàn)低溫可以促進(jìn)其中的果蠅胰島素RNA的合成和胰島素的分泌。說明IPCs做為一個(gè)發(fā)育調(diào)控關(guān)鍵細(xì)胞和一個(gè)低溫興奮性神經(jīng)元在低溫促進(jìn)幼蟲發(fā)育過程中的重要性。更進(jìn)一步,我們通過篩選得到一個(gè)標(biāo)記果蠅頭部神經(jīng)元的11216-Gal4品系,在體鈣成像實(shí)驗(yàn)的數(shù)據(jù)表明,低溫刺激可以激活11216-Gal4標(biāo)記的神經(jīng)元,通過鑒定和比對(duì),我們確認(rèn)這些神經(jīng)元為幼蟲的初級(jí)冷覺感知神經(jīng)元。我們利用GRASP (GFP reconstitution across synaptic partners)技術(shù),證明這些11216-Gal4標(biāo)記的初級(jí)冷覺感知神經(jīng)元的軸突與IPCs的樹突形成物理上的突觸聯(lián)系；利用光遺傳結(jié)合鈣成像的方法,我們發(fā)現(xiàn)激活11216-Gal4標(biāo)記的初級(jí)冷覺感知神經(jīng)元可以激活I(lǐng)PCs,說明這兩群神經(jīng)元形成功能性的神經(jīng)環(huán)路。同時(shí)我們發(fā)現(xiàn),激活11216-Gal4標(biāo)記的初級(jí)冷覺感知神經(jīng)元可以促進(jìn)IPCs中的果蠅胰島素RNA的合成和胰島素的分泌,并且導(dǎo)致果蠅的體型增大,很好的再現(xiàn)了低溫刺激的效果。我們的研究揭示了冷覺信號(hào)刺激胰島素合成分泌并促進(jìn)個(gè)體生長(zhǎng)的神經(jīng)信號(hào)傳遞通路,為TSR提供了一個(gè)神經(jīng)環(huán)路水平的機(jī)制性解釋,也為Bergmann's rule的解釋提供了借鑒。
[Abstract]:The environmental factor is a research topic of enduring influence on individual development, among other factors, climate change, temperature change is everywhere, it has great influence on the.19 century animal ontogeny and population evolution, the German biologist Carl Bergmann discovered that size and survival latitude is positively related to the individual animal the temperature is, the higher the latitude, the cold area, the individual animal bigger in lower latitude area, temperature is warm, the smaller animal individual. This is the famous Bergmann's rule, its general rule summarizes the influence of temperature on the survival of warm blooded animal size. In vertebrates, the animal body with temperature. The negative correlation is more prevalent, we call the body temperature regulation (Temperature-Size Rule, TSR). From the beginning of nineteenth Century, people study and investigation of all kinds of animal, try Explain the temperature influence reasons of animal body, so far we on the molecular and cellular mechanisms are still poorly understood. Animal model of Drosophila is a good carrier to study the influence of temperature size rule, in early 1940, people have begun to use the influence of temperature on size of fruit flies, flies its survival in the suitable temperature range, the size of mature body the temperature has a strong plasticity, such as low temperature (18 DEG C) were smaller than Drosophila under high temperature (25 DEG C) about feeding 10%. Fruit fly Drosophila larvae stage of development is accumulated and decisive stage of mature body size, so we mainly study the developmental mechanism of larvae of Drosophila larvae. And as adults, with rich perception system, including chemical sensing, mechanical sensing, temperature sensing. In the primary distribution of peripheral larvae to perceive God rich by yuan, their investment Shot to the brain central neurons form neural network is simple and functional. The primary sensory neurons is animal perception of external environmental factors (temperature) of the temperature sensing we the only way which must be passed, nervous system of Drosophila larvae as the starting point, the excitability of neurons in the artificial control means, combined with the basic research of cellular and molecular mechanism of temperature effect detection method the ontogeny of Drosophila. Insulin is one of the key molecules regulating larval development and species conservation, several major Drosophila insulin producing cells by insulin IPCs (insulin producing cells), a secretory neuron, synthesis and secretion. We found that blocking the IPCs can eliminate the effect of low temperature effects of larval size increased, while the activation of IPCs can be to promote the increase in the size of larvae, the imaging experiment of body calcium, we found that low temperature stimulation can be excited by IPCs. Light quantitative PCR, immunohistochemistry and Western blotting experiments, we found that low temperature can promote the synthesis and secretion of insulin in the Drosophila insulin RNA. IPCs as a key developmental control cell and a low temperature low temperature promote excitatory neurons in the larval developmental importance process. Further, we obtained through screening the 11216-Gal4 strain of a Drosophila head marker of neurons, that in body calcium imaging experiment data, low temperature stimulation can activate 11216-Gal4 labeled neurons, through the identification and comparison, we confirmed that these neurons of primary sensory neurons of larvae. We use GRASP (GFP reconstitution across synaptic partners) technology, that axons and the primary IPCs cryesthesia sensory neurons of these 11216-Gal4 markers for dendritic synapses physically; using light calcium combined with genetic The imaging method, we found that the initial activation of 11216-Gal4 labeled neurons cold perception can activate IPCs, indicating that the two groups of neurons form functional neural circuits. At the same time, we found that the primary sensory neurons labeled by 11216-Gal4 activation of IPCs can promote the synthesis and secretion of insulin in the fruit fly RNA insulin, and increase the size of Drosophila melanogaster well, the reproduction of the low temperature stimulation effect. Our study reveals the cold stimulation of insulin synthesis and secretion and promote individual growth of neural signal pathway, provides a mechanism of a neural circuit level explanation for TSR, but also provide a reference for the interpretation of rule Bergmann's.

【學(xué)位授予單位】：浙江大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2016
【分類號(hào)】：Q43

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