中华齿刺甲的生物学特性及其抗冻蛋白的功能研究
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摘要
中华齿刺甲(Oodescelis chinensis Kaszab)隶属于鞘翅目，拟步甲科，刺甲族，为中国特有种昆虫，仅分布于我国新疆的荒漠和半荒漠地区。中华齿刺甲没有荒漠拟步甲普遍具有的自相残杀习性，具有明显的雌雄二型性特征，更适合作模式生物。中华齿刺甲的越冬地在土壤表面，虫体在低于过冷却点的温度下仍能存活，为耐冻昆虫，是研究耐冻昆虫抗冻蛋白的良好材料。在此之前，该昆虫还没有在室内饲养成功，尚不清楚该昆虫的生物学特性，对于其雌雄二型性特征也缺乏了解，更不知晓耐冻昆虫中抗冻蛋白的功能。本研究建立了有效饲养中国特有种拟步甲科昆虫中华齿刺甲的方法，对其生物学特性、雌雄二型性和雌雄鉴别进行了研究，同时克隆得到了11个抗冻蛋白基因，并对其中的3种抗冻蛋白进行了结构和功能测试，为该荒漠昆虫的后续深入研究奠定了基础。
采用加有湿润棉花的玻璃培养皿孵育卵、预蛹、蛹和初孵成虫。用含有湿润沙土的昆虫饲养管饲养幼虫，由于毛细作用，饲养管中沙子的含水量由下向上逐渐降低，表面为干沙层，在其上面加麦麸饲养幼虫。结果表明，此方法能使昆虫完成其正常的生长发育和繁殖过程，其卵、1龄幼虫、2龄以后幼虫、预蛹、蛹和幼嫩成虫的存活率分别为86.39±4.43％、92.85±1.22、55.22±5.65％、82.35±4.37、85.13±3.41％和86.00±7.37％。此方法能使昆虫在室内顺利完成其生活史，并能正常交配，繁殖后代。
中华齿刺甲为夜行性昆虫，在乌鲁木齐1年可以完成1代，有世代重叠现象，成虫性比约为1:1。雌性成虫的体重和鞘翅宽显著大于雄虫。幼虫共15龄，无自相残杀的习性。在昆虫的每个虫态，都有昆虫某些部位体色加深的变化，一般从乳白到黄色，再到红褐或者黑色。成虫产卵期约为4个月，平均单雌产卵量为1032枚。雌雄性成虫和蛹有明显的性别二型性特征。
中华齿刺甲成虫有明显的3个外部可见的雌雄二型性特征和1个第8腹板上的性别鉴别特征。列举如下。第一，雄性成虫的前足和中足跗节有宽的爪垫，利用扫描电镜对爪垫进行观察的结果显示，爪垫是由一个个纤细的扁平条状结构组成，前段稍尖并向背面弯曲，背面有竹节內隔状螺纹。第二，雄性成虫的后足腹面覆盖有一层刚毛刷。第三，雄虫腹面的第1、2、5腹节各有1个由刚毛组成的圆形刚毛垫。第四，雄性的第8腹板尖端有1个三角形的缺刻。以上特征在雌性成虫中缺无。第4个特征在小胸鳖甲(Microderapunctipennis)、细胸鳖甲(Colposcelis microderoides microderoides)、光滑鳖甲(Anatolicapolita borealis)、异长足漠甲(Adesmia anomala dejeani)、苏氏漠甲(Sternoplax souvorowiana)和伪东鳖甲(Anatolica pseudiduma)等拟步甲昆虫中普遍存在，只存在颜色上的细微差别。基于该特征开发出了一种性别鉴别方法——―第8腹板探测法‖，来鉴别荒漠拟步甲成虫的雌雄，该方法可以被广泛用于拟步甲成虫的雌雄快速无损鉴别。中华齿刺甲蛹也有3个明显的雌雄二型性特征。第一，类似于雄虫，雄蛹的前足和中足跗节也明显宽于雌性，将来发育成爪垫。第二，雄蛹的第8腹节上没有明显的生殖孔。第三，雄蛹第8腹节窄小，末端上有2个紧邻的突起；雌蛹第8腹节宽大，末端两侧有2个分离的乳头状突起。其中，第2和第3个特征在拟步甲科昆虫的蛹中具有普遍性，可以用于其他拟步甲昆虫蛹的无损雌雄鉴别。中华齿刺甲成虫跗爪上的爪垫和第8腹板上的缺刻等特征，在雌雄性成虫抱对时，可能有利于雄虫牢固抓住雌虫，有利于交配的顺利进行。
从耐冻昆虫中华齿刺甲的越冬成虫中克隆得到了11个抗冻蛋白基因，在成熟的蛋白质序列中，一级结构由l2个氨基酸TCT(I)xSxxCxxAx (X表示其它的任意氨基酸残基)的TCT(I)重复序列组成，每隔6个氨基酸残基，就有一个半胱氨酸残基重复。这些蛋白前端均含有信号肽，信号肽长23个氨基酸。这11种抗冻蛋白基因主要分为两大类型，第一类，编码一级结构由TCTxSxxCxxAx (TCT重复序列)组成的抗冻蛋白基因，再根据其氨基酸个数继续分为编码122个氨基酸、95(94)个氨基酸和83个氨基酸的3种抗冻蛋白基因。第二类，编码一级结构由TCIxSxxCxxAx (TCI重复序列)组成的抗冻蛋白基因，该类只有1种基因，为编码97个氨基酸的抗冻蛋白基因Ocafp2。本研究从中选择了3种抗冻蛋白基因Ocafp1-3(编码122个氨基酸)、Ocafp2(编码97个氨基酸)和Ocafp4(编码95个氨基酸)，构建到不同的原核表达质粒(pET28a-egfp和pET32a)上，通过大肠杆菌(Escherichia coli BL21)(DE3)实现了可溶性表达，经过镍柱纯化获得了带有有不同标签蛋白的重组抗冻蛋白EGFP-OcAFPs和TrxA-OcAFPs。并对其进行了结构和功能的测试，主要研究结果如下：用圆二色谱仪测定其二级结构的结果表明，3种中华齿刺甲的抗冻蛋白都以β-sheet为主；差示扫描量热仪的测定结果显示，OcAFP1-3和OcAFP4具有中度的热滞活性(THA)和酸碱稳定性，但是不耐受60℃以上的高温，OcAFP2的热滞活性低至几乎测不出来；热重分析结果显示OcAFP4的亲水性强于OcAFP1-3，OcAFP2亲水性最弱；纳升渗透压仪观测结果显示，OcAFP1-3和OcAFP4修饰冰晶分别呈六边形和四边形，OcAFP2则呈现骨针形；OcAFP1-3和OcAFP4使冰晶沿着a轴生长，OcAFP2则使冰晶沿着c轴生长。这预示着OcAFP1-3和OcAFP4主要结合到了冰晶的基面，显示其还具有高活性抗冻蛋白的特性；而OcAFP2主要结合到了冰晶的棱面，显示其还具有中度活性抗冻蛋白的特性。以上热滞值和冰晶形态结果，说明耐冻昆虫中华齿刺甲的抗冻蛋白(OcAFP1-3和OcAFP4)兼有避冻昆虫和鱼类抗冻蛋白的双重活性，OcAFP2兼有耐冻植物抗冻蛋白和鱼类抗冻蛋白的双重活性。这些新发现，弥补了对于耐冻昆虫体内抗冻蛋白研究方面的空缺，增进了我们对抗冻蛋白的认知。
Oodescelis chinensis Kaszab (Coleoptera: Tenebrionidae) is a Chinese endemic species,which is only distributed in desert and semidesert regions of Xinjiang, China. Oodescelischinensis possesses significant ecological and economic values, and is suitable to be used as amodel creature because of its obviously sexual dimorphism and the habit of not killing eachother. O. chinensis lives through winter on the surface of soil, and the insect can surviveafter supercooling point test experiment. It is a suitable creature for studying antifreezeproteins. But its biology and antifreeze proteins remain largely unknown. Also, its laboratoryrearing and its sexual dimorphism are yet to be determined. In this study, the laboratoryrearing methods, the biological characteristics and the sexing methods of O. chinensis wereexamined. Furthermore, eleven antifreeze protein genes were isolated from overwinter adults,and the structures and functions of three antifreeze proteins were studied. This study laidfoundation for further research of the desert insect.
The Glass Petri dishes added with humid cotton were used to cultivate the insect eggs,prepupae, pupae and teneral adults. The larvae were reared in insect rearing tubes with moistdesert sand. The water content of desert sand decreased from bottom to upper part because ofcapillarity, and the surface of the desert soil was dry. The wheat bran was added to raise larvaeof O. chinensis. It can finish its life cycle under the rearing condition. The survival rates ofeggs, first instar larvae, larvae after second instar, prepupae, pupae, and teneral adults were86.39±4.43％,92.85±1.22,55.22±5.65％,82.35±4.37,85.13±3.41％and86.00±7.37％,respectively. Oodescelis chinensis can successfully finish its life cycle, and the insect can passto the next generation normally in laboratory.
O. chinensis adopts nocturnality to survive desert environment. It had one-year life cycleand overlapping generation, in Urumqi. The female/male ratio of adult insect was1:1. Theweight and elytrum length of female was obviously greater than that of male in adult. Therewere15instars during larvae stage. The coloration of each developmental stage gradually changed from creamy white to light brownish, brunneus or black. The egg laying period offemale adult lasts4months. On average, each female produced1032eggs. The adults andpupae have abviously sexual dimorphism apparences.
The adult of Oodescelis chinensis has three externally visible gender-specificcharacteristics, and an inner characteristic on the8thsternite for sex determination. First, thereare broad claw pads in tarsus of the male adult fore legs and middle legs. These claw padswere consisted of slim flat bands which can be seen with scanning electron microscope. Theforeparts of the slim flat bands are slightly sharp and the tips are bent back. The back of thestructure has bamboo-shaped screw threads. Second, the hind legs of male adult are coveredwith bristle layers. Third, the first, second and fifth abdominal segments of male adult eachhas one bristle pad. The fourth, there is a trilateral indentation on the8thabdominal sternite inmale. However, these characteristics in male are absent in female adult. The fouthcharacteristic is very common in tenebrionidae beetles, such as Microdera punctipennis,Colposcelis microderoides microderoides, Anatolica polita borealis, Adesmia anomaladejeani, Sternoplax souvorowiana and Anatolica pseudiduma. A new method based on thedifference of the8thsternite was developed to determinate the sexes of desert beelte adults.This method could be widely used in sexing desert beeltes with100%accuracy and withoutdemage. The pupaes of O. chinensis also has three sexual characteristics. First, similar to maleadults, the tarsus of the male pupal fore legs and middle legs are obviously wider than that offemale, which will be developed to claw pads. Second, there are no obviously genitalopenings on the8thabdominal sternite in male pupae. Third, the8thabdominal sternite of malepupae is narrower than that of female. There are two closed prominencies at the end of the8thabdominal sternite in male pupae, but the two prominencies of female pupae are separated andlooked like nipples. The second and the third characteristic of pupae are universal inTenebrionidae beetle pupae, so it can be used in pupal sex determination of tenebrionidi beetlewithout damage. The claw pads in tarsus and indentation on the8thabdominal sternite of Oodescelis chinensis male adults are helpful during coupling and mating.
Eleven antifreeze genes were isolated from antifreeze tolerant desert beetle O. chinensisin this research. The primary structure of these protein comprised of tandem repeats of12-aasequence [TCT(I)xSxxCxxAx] with regularly spaced Cys at intervals of6-aa residues, andthere are23-aa signal peptides in the N-terminal part of these antifreeze proteins. The genesof these antifreeze proteins can be divided into two types. One is the antifreeze genes thatcode primary structure of TCT(I)xSxxCxxAx repeated sequence. These type genes includecoding122-aa,94(95)-aa and83-aa antifreeze protein genes. The other is the gene that codeprimary structure of TCIxSxxCxxAx repeated sequence, which only codes97-aa. Theantifreeze genes Ocafp1-3(122-aa), Ocafp2(97-aa) and Ocafp4(95-aa) were selected toconstructe with different expressive plasmids (pET28a-egfp and pET32a), and thecorresponding mature peptide has been expressed in E. coli, respectively. The recombinantantifreeze proteins EGFP-OcAFPs and TrxA-OcAFPs were obtained after purification byNi-IDA. The structions and functions of these recombination antifreeze proteins are asfollows. The secondary structures of these antifreeze proteins are mainly β-sheet which isconfirmed by circular dichroism spectrometer. The results of differential scanning calorimetryshowed that OcAFP1-3and OcAFP4has moderate thermal hysteresis activity (THA) and pHand heat stability, but they can‘t endure tempersture higher than60℃. The THAof OcAFP2istoo low to be measured. TGA themogravimetric analysis suggests that the hydrophilicity ofOcAFP4is stronger than that of OcAFP1-3, and the hydrophilicity of OcAFP2is the weakestamong these antifreeze proteins. The results observed with nanolitre osmometer showed thatthe modification ice crystal of OcAFP1-3, OcAFP4and OcAFP2are hexagon, quadrilateraland bone nails shape. OcAFP1-3and OcAFP4made ice crystal growing along a-axis, andOcAFP2made ice crystal growing along c-axis. These phenomena indicate that OcAFP1-3and OcAFP4are mainly combined to the basal plate of the ice crystal, and OcAFP2is mainlycombinated to the prism faces of the ice crystal. Our results indicate that OcAFP1-3and OcAFP4have both characteristics of antifreeze protein with high and moderate activities;while OcAFP2has characteristics of antifreeze protein with moderate and low activities.These new discoveries shed light on our understanding of the antifreeze proteins obtainedfrom freeze tolerant insect.
采用加有湿润棉花的玻璃培养皿孵育卵、预蛹、蛹和初孵成虫。用含有湿润沙土的昆虫饲养管饲养幼虫，由于毛细作用，饲养管中沙子的含水量由下向上逐渐降低，表面为干沙层，在其上面加麦麸饲养幼虫。结果表明，此方法能使昆虫完成其正常的生长发育和繁殖过程，其卵、1龄幼虫、2龄以后幼虫、预蛹、蛹和幼嫩成虫的存活率分别为86.39±4.43％、92.85±1.22、55.22±5.65％、82.35±4.37、85.13±3.41％和86.00±7.37％。此方法能使昆虫在室内顺利完成其生活史，并能正常交配，繁殖后代。
中华齿刺甲为夜行性昆虫，在乌鲁木齐1年可以完成1代，有世代重叠现象，成虫性比约为1:1。雌性成虫的体重和鞘翅宽显著大于雄虫。幼虫共15龄，无自相残杀的习性。在昆虫的每个虫态，都有昆虫某些部位体色加深的变化，一般从乳白到黄色，再到红褐或者黑色。成虫产卵期约为4个月，平均单雌产卵量为1032枚。雌雄性成虫和蛹有明显的性别二型性特征。
中华齿刺甲成虫有明显的3个外部可见的雌雄二型性特征和1个第8腹板上的性别鉴别特征。列举如下。第一，雄性成虫的前足和中足跗节有宽的爪垫，利用扫描电镜对爪垫进行观察的结果显示，爪垫是由一个个纤细的扁平条状结构组成，前段稍尖并向背面弯曲，背面有竹节內隔状螺纹。第二，雄性成虫的后足腹面覆盖有一层刚毛刷。第三，雄虫腹面的第1、2、5腹节各有1个由刚毛组成的圆形刚毛垫。第四，雄性的第8腹板尖端有1个三角形的缺刻。以上特征在雌性成虫中缺无。第4个特征在小胸鳖甲(Microderapunctipennis)、细胸鳖甲(Colposcelis microderoides microderoides)、光滑鳖甲(Anatolicapolita borealis)、异长足漠甲(Adesmia anomala dejeani)、苏氏漠甲(Sternoplax souvorowiana)和伪东鳖甲(Anatolica pseudiduma)等拟步甲昆虫中普遍存在，只存在颜色上的细微差别。基于该特征开发出了一种性别鉴别方法——―第8腹板探测法‖，来鉴别荒漠拟步甲成虫的雌雄，该方法可以被广泛用于拟步甲成虫的雌雄快速无损鉴别。中华齿刺甲蛹也有3个明显的雌雄二型性特征。第一，类似于雄虫，雄蛹的前足和中足跗节也明显宽于雌性，将来发育成爪垫。第二，雄蛹的第8腹节上没有明显的生殖孔。第三，雄蛹第8腹节窄小，末端上有2个紧邻的突起；雌蛹第8腹节宽大，末端两侧有2个分离的乳头状突起。其中，第2和第3个特征在拟步甲科昆虫的蛹中具有普遍性，可以用于其他拟步甲昆虫蛹的无损雌雄鉴别。中华齿刺甲成虫跗爪上的爪垫和第8腹板上的缺刻等特征，在雌雄性成虫抱对时，可能有利于雄虫牢固抓住雌虫，有利于交配的顺利进行。
从耐冻昆虫中华齿刺甲的越冬成虫中克隆得到了11个抗冻蛋白基因，在成熟的蛋白质序列中，一级结构由l2个氨基酸TCT(I)xSxxCxxAx (X表示其它的任意氨基酸残基)的TCT(I)重复序列组成，每隔6个氨基酸残基，就有一个半胱氨酸残基重复。这些蛋白前端均含有信号肽，信号肽长23个氨基酸。这11种抗冻蛋白基因主要分为两大类型，第一类，编码一级结构由TCTxSxxCxxAx (TCT重复序列)组成的抗冻蛋白基因，再根据其氨基酸个数继续分为编码122个氨基酸、95(94)个氨基酸和83个氨基酸的3种抗冻蛋白基因。第二类，编码一级结构由TCIxSxxCxxAx (TCI重复序列)组成的抗冻蛋白基因，该类只有1种基因，为编码97个氨基酸的抗冻蛋白基因Ocafp2。本研究从中选择了3种抗冻蛋白基因Ocafp1-3(编码122个氨基酸)、Ocafp2(编码97个氨基酸)和Ocafp4(编码95个氨基酸)，构建到不同的原核表达质粒(pET28a-egfp和pET32a)上，通过大肠杆菌(Escherichia coli BL21)(DE3)实现了可溶性表达，经过镍柱纯化获得了带有有不同标签蛋白的重组抗冻蛋白EGFP-OcAFPs和TrxA-OcAFPs。并对其进行了结构和功能的测试，主要研究结果如下：用圆二色谱仪测定其二级结构的结果表明，3种中华齿刺甲的抗冻蛋白都以β-sheet为主；差示扫描量热仪的测定结果显示，OcAFP1-3和OcAFP4具有中度的热滞活性(THA)和酸碱稳定性，但是不耐受60℃以上的高温，OcAFP2的热滞活性低至几乎测不出来；热重分析结果显示OcAFP4的亲水性强于OcAFP1-3，OcAFP2亲水性最弱；纳升渗透压仪观测结果显示，OcAFP1-3和OcAFP4修饰冰晶分别呈六边形和四边形，OcAFP2则呈现骨针形；OcAFP1-3和OcAFP4使冰晶沿着a轴生长，OcAFP2则使冰晶沿着c轴生长。这预示着OcAFP1-3和OcAFP4主要结合到了冰晶的基面，显示其还具有高活性抗冻蛋白的特性；而OcAFP2主要结合到了冰晶的棱面，显示其还具有中度活性抗冻蛋白的特性。以上热滞值和冰晶形态结果，说明耐冻昆虫中华齿刺甲的抗冻蛋白(OcAFP1-3和OcAFP4)兼有避冻昆虫和鱼类抗冻蛋白的双重活性，OcAFP2兼有耐冻植物抗冻蛋白和鱼类抗冻蛋白的双重活性。这些新发现，弥补了对于耐冻昆虫体内抗冻蛋白研究方面的空缺，增进了我们对抗冻蛋白的认知。
Oodescelis chinensis Kaszab (Coleoptera: Tenebrionidae) is a Chinese endemic species,which is only distributed in desert and semidesert regions of Xinjiang, China. Oodescelischinensis possesses significant ecological and economic values, and is suitable to be used as amodel creature because of its obviously sexual dimorphism and the habit of not killing eachother. O. chinensis lives through winter on the surface of soil, and the insect can surviveafter supercooling point test experiment. It is a suitable creature for studying antifreezeproteins. But its biology and antifreeze proteins remain largely unknown. Also, its laboratoryrearing and its sexual dimorphism are yet to be determined. In this study, the laboratoryrearing methods, the biological characteristics and the sexing methods of O. chinensis wereexamined. Furthermore, eleven antifreeze protein genes were isolated from overwinter adults,and the structures and functions of three antifreeze proteins were studied. This study laidfoundation for further research of the desert insect.
The Glass Petri dishes added with humid cotton were used to cultivate the insect eggs,prepupae, pupae and teneral adults. The larvae were reared in insect rearing tubes with moistdesert sand. The water content of desert sand decreased from bottom to upper part because ofcapillarity, and the surface of the desert soil was dry. The wheat bran was added to raise larvaeof O. chinensis. It can finish its life cycle under the rearing condition. The survival rates ofeggs, first instar larvae, larvae after second instar, prepupae, pupae, and teneral adults were86.39±4.43％,92.85±1.22,55.22±5.65％,82.35±4.37,85.13±3.41％and86.00±7.37％,respectively. Oodescelis chinensis can successfully finish its life cycle, and the insect can passto the next generation normally in laboratory.
O. chinensis adopts nocturnality to survive desert environment. It had one-year life cycleand overlapping generation, in Urumqi. The female/male ratio of adult insect was1:1. Theweight and elytrum length of female was obviously greater than that of male in adult. Therewere15instars during larvae stage. The coloration of each developmental stage gradually changed from creamy white to light brownish, brunneus or black. The egg laying period offemale adult lasts4months. On average, each female produced1032eggs. The adults andpupae have abviously sexual dimorphism apparences.
The adult of Oodescelis chinensis has three externally visible gender-specificcharacteristics, and an inner characteristic on the8thsternite for sex determination. First, thereare broad claw pads in tarsus of the male adult fore legs and middle legs. These claw padswere consisted of slim flat bands which can be seen with scanning electron microscope. Theforeparts of the slim flat bands are slightly sharp and the tips are bent back. The back of thestructure has bamboo-shaped screw threads. Second, the hind legs of male adult are coveredwith bristle layers. Third, the first, second and fifth abdominal segments of male adult eachhas one bristle pad. The fourth, there is a trilateral indentation on the8thabdominal sternite inmale. However, these characteristics in male are absent in female adult. The fouthcharacteristic is very common in tenebrionidae beetles, such as Microdera punctipennis,Colposcelis microderoides microderoides, Anatolica polita borealis, Adesmia anomaladejeani, Sternoplax souvorowiana and Anatolica pseudiduma. A new method based on thedifference of the8thsternite was developed to determinate the sexes of desert beelte adults.This method could be widely used in sexing desert beeltes with100%accuracy and withoutdemage. The pupaes of O. chinensis also has three sexual characteristics. First, similar to maleadults, the tarsus of the male pupal fore legs and middle legs are obviously wider than that offemale, which will be developed to claw pads. Second, there are no obviously genitalopenings on the8thabdominal sternite in male pupae. Third, the8thabdominal sternite of malepupae is narrower than that of female. There are two closed prominencies at the end of the8thabdominal sternite in male pupae, but the two prominencies of female pupae are separated andlooked like nipples. The second and the third characteristic of pupae are universal inTenebrionidae beetle pupae, so it can be used in pupal sex determination of tenebrionidi beetlewithout damage. The claw pads in tarsus and indentation on the8thabdominal sternite of Oodescelis chinensis male adults are helpful during coupling and mating.
Eleven antifreeze genes were isolated from antifreeze tolerant desert beetle O. chinensisin this research. The primary structure of these protein comprised of tandem repeats of12-aasequence [TCT(I)xSxxCxxAx] with regularly spaced Cys at intervals of6-aa residues, andthere are23-aa signal peptides in the N-terminal part of these antifreeze proteins. The genesof these antifreeze proteins can be divided into two types. One is the antifreeze genes thatcode primary structure of TCT(I)xSxxCxxAx repeated sequence. These type genes includecoding122-aa,94(95)-aa and83-aa antifreeze protein genes. The other is the gene that codeprimary structure of TCIxSxxCxxAx repeated sequence, which only codes97-aa. Theantifreeze genes Ocafp1-3(122-aa), Ocafp2(97-aa) and Ocafp4(95-aa) were selected toconstructe with different expressive plasmids (pET28a-egfp and pET32a), and thecorresponding mature peptide has been expressed in E. coli, respectively. The recombinantantifreeze proteins EGFP-OcAFPs and TrxA-OcAFPs were obtained after purification byNi-IDA. The structions and functions of these recombination antifreeze proteins are asfollows. The secondary structures of these antifreeze proteins are mainly β-sheet which isconfirmed by circular dichroism spectrometer. The results of differential scanning calorimetryshowed that OcAFP1-3and OcAFP4has moderate thermal hysteresis activity (THA) and pHand heat stability, but they can‘t endure tempersture higher than60℃. The THAof OcAFP2istoo low to be measured. TGA themogravimetric analysis suggests that the hydrophilicity ofOcAFP4is stronger than that of OcAFP1-3, and the hydrophilicity of OcAFP2is the weakestamong these antifreeze proteins. The results observed with nanolitre osmometer showed thatthe modification ice crystal of OcAFP1-3, OcAFP4and OcAFP2are hexagon, quadrilateraland bone nails shape. OcAFP1-3and OcAFP4made ice crystal growing along a-axis, andOcAFP2made ice crystal growing along c-axis. These phenomena indicate that OcAFP1-3and OcAFP4are mainly combined to the basal plate of the ice crystal, and OcAFP2is mainlycombinated to the prism faces of the ice crystal. Our results indicate that OcAFP1-3and OcAFP4have both characteristics of antifreeze protein with high and moderate activities;while OcAFP2has characteristics of antifreeze protein with moderate and low activities.These new discoveries shed light on our understanding of the antifreeze proteins obtainedfrom freeze tolerant insect.
引文
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