红螯光壳螯虾亲虾脂类营养需求及脂肪酸结合蛋白的功能研究
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摘要
本研究采用水生动物营养学、生物化学、分子生物学和生物光子学等研究手段,较为系统和深入的研究了不同脂肪源饲料对雌性红螯光壳螯虾相关营养生理学参数的影响,并在此基础上从饲料的胆固醇添加量入手,研究了胆固醇在提高其生长和生殖性能方面的作用,初步阐明了脂类营养对红螯光壳螯虾生长和生殖的调节功能。同时有针对性地利用分子生物学和生物光子学对与红螯光壳螯虾脂类摄取和代谢有关的脂肪酸结合蛋白FABP进行深入研究,以期阐明脂肪对其生长和生殖调控的机理。研究成果不仅对红螯光壳螯虾的亲体培育及优化红螯光壳螯虾人工饲料的配置具有重要的现实指导意义,也为虾蟹类营养学调控的深入研究提供理论依据。
1.不同脂肪源饲料对雌性红鳌光壳螯虾生长及组织生化组成的影响
取平均体重为19.25±0.72g雌性红螯光壳螯虾,以鱼油、花生油、豆油和猪油作为脂肪源配制4种粗蛋白含量为33%,粗脂肪为8.5%的饲料,并以商业饲料作为对照投喂60天,研究不同脂肪源条件对雌性红螯光壳螯虾亲虾生长及组织生化组成的影响。结果显示：不同脂肪源饲料对雌性红螯光壳螯虾亲虾的增重、特殊生长率、存活率、饲料系数均有显著影响,而在本实验中对GSI无显著影响。其中以豆油作为脂肪源的饲料投喂的亲虾在增重、存活率等指标上均显著优于其他各组。在基本生化组成上,除了花生油组亲虾在肌肉和肝胰腺中脂肪和蛋白含量上略低外,其余各组无显著差异。在脂肪酸组成上,肌肉中主要的脂肪酸为c16:0, c18:0, c18:1n9, c18:2n6 and c20:5n3,在肝胰腺中为c16：0,c18：ln9,c18：2n6,肌肉中的c20:5n-3 (EPA)含量显著高于肝胰腺。不同饲料组对肝胰腺中的脂肪酸组成影响非常显著,饲料中SFA/MUFA/PUFA的比例直接影响了肝胰腺中脂肪酸的组成。各组肌肉脂肪酸的SFA/PUFA占总脂的比率没有显著的差异,仅MUFA有一定差异,其中第5组的的MUFA含量在总脂中所占比例最小。结果显示,在粗蛋白含量为33%,粗脂肪为8.5%条件下采用豆油作为脂肪源较适于此阶段的雌性红螯光壳螯虾生长。
2.不同脂肪源饲料对雌性红鳌光壳螯虾消化酶活力以及Vg基因表达的影响
在以不同脂肪源饲料投喂60天后,通过对雌性红螯光壳螯虾亲虾肝胰腺中消化酶活力以及卵黄蛋白源Vg基因的表达量的检测,研究不同脂肪源饲料对雌性亲虾的影响。结果显示消化酶活力受到不同脂肪源的影响,其中鱼油组、豆油组和商业饲料组脂肪酶活力显著高于花生油组和猪油组,但三者间无显著性差异；胰蛋白酶则以鱼油组、花生油组和豆油组较高,猪油组和商业饲料组相对较底；不同脂肪源饲料对淀粉酶活力影响相对较小,除花生油组略高外,其余各组无显著性差异,在纤维素酶活力上,除猪油组显著较低外,其余各组无显著差异。总体上讲,豆油组各消化酶活力均较好。虽然不同脂肪源组亲虾在GSI上没有显著差异,在肝胰腺Vg表达量上则有显著差异。其中豆油组投喂的亲虾最高的Vg表达量暗示了豆油较好的促卵巢成熟效果。结合生长指、消化酶活力和Vg表达,结果显示在本实验条件下以豆油作为脂肪源较适于雌性亲虾生长及生殖。
3.不同胆固醇含量饲料对红鳌光壳螫虾雌虾生长及生殖的影响
取平均体重为19.53+1.03g雌性红螯光壳螯虾,以0%、0.25%、0.5%、0.75%、1%胆固醇添加量配置5种粗蛋白含量为34%,粗脂肪为8.5%的饲料投喂60天。通过对不同实验组雌性红螯光壳螯虾亲虾的基本生长指标、脂肪酸组成、消化酶活力以及Vg表达量的研究,通过非线性回归来寻找此阶段雌性红螯光壳螯虾的胆固醇最适添加量。结果显示不同胆固醇添加量对肌肉和肝胰腺脂肪酸组成影响不显著,但对其基本生长指标、消化酶活力以及Vg表达均有显著影响。其中未添加组在增重、存活率等生长指标,脂肪酶、胰蛋白酶活力以及Vg表达量上均比添加胆固醇组低。在添加胆固醇的组别中,0.25%及0.5%的胆固醇添加组亲虾在增重率、SGR、存活率等生长指标,脂肪酶、胰蛋白酶活力以及Vg表达量上均显示出较好的效果,而高添加量组(1%)虽然在Vg表达量上最高,但在其他指标上则显示出对生长的抑制作用。对增重、SGR、存活率、FCR以及脂肪酶和胰蛋白酶活力的回归分析结果显示,此阶段雌虾胆固醇最适添加量为0.55%。因此,在雌性红螯光壳螯虾卵黄发生的中后期,可将胆固醇添加量增至0.75%以满足其卵巢快速发育期需求。
4.红螯光壳螯虾Cq-FABP基因的克隆和序列特征分析
通过兼并引物PCR获得部分片段后,采用RACE技术成功的克隆了红螯光壳螯虾脂肪酸结合蛋白FABP基因的全长cDNA序列。该序列全长772bp,包含一个82bp的5’-UTR区域和一个291bp的3’-UTR区域,开放阅读框长399bp,编码132个氨基酸。通过生物信息学方法对其二级、三级结构预测结果显示该蛋白具有FABP典型的特征：10个β折叠和2个α螺旋,并通过β折叠和α螺旋形成能够容纳疏水配基的桶装结构。同源性比对结果显示红螯光壳螯虾脂肪酸结合蛋白Cq-FABP与通讯螯虾具有最高的同源性,为90.9%；与其他已知的甲壳类十足目种类的同源性则相对较低,和中华绒螯蟹、克氏原螯虾、刀额新对虾、凡纳滨对虾以及斑节对虾FABP的同源性分别为56.8%、56.8%、53.8%、55.3%和53.8%。而与人类多种FABP的比对结果显示Cq-FABP与人类脑型脂肪酸结合蛋白B-FABP具有最高的同源性,为40.9%,与心型和肌肉型FABP的同源性次之,为38.6%,这也暗示了其可能具有功能的多样性。系统进化结果也显示红螯光壳螯虾FABP虽然与其他甲壳类十足日FABP同属于一簇,但其与通讯螯虾同属与一个分支,其他十足目种类则属于另一分支。
5.红螯光壳螯虾Cq-FABP基因的表达特征分析
采用实时定量PCR技术研究了红螯光壳螯虾Cq-FABP基因在不同组织中的表达以及受不同脂肪源饲料影响后在肝胰腺中的表达量变化,均以β-actin作为内参。结果发现虽然在多个组织中均检测到Cq-FABP表达,但主要在肝胰腺组织中表达。在肠中也有少量表达,但表达量仅为肝胰腺腺中的1/70,在卵巢、触角腺、血淋巴、腮、胃和精巢中也有极微量的表达,但在肌肉中几乎无表达。由于其表达量最高的在肝胰腺组织,其次在肠,暗示了Cq-FABP的功能可能主要涉及脂肪酸的摄取和利用。以不同脂肪源饲料投喂的亲虾肝胰腺中Cq-FABP的表达量与脂肪源中的长链脂肪酸比例呈正比,但长链脂肪酸比例较为接近的花生油和豆油组表达量却较低。根据豆油组的亲虾的生长情况及的Vg高表达量推测可能与Cq-FABP的表达有关,而Cq-FABP与人B-FABP的最高同源性也暗示了不同的脂肪酸可能可以通过调节Cq-FABP的方式参与到信号调控中。
6. Cq-FABP蛋白的表达纯化及与脂肪酸结合特性的研究
将红螯光壳螯虾脂肪酸结合蛋白Cq-FABP基因重组入原核表达载体pET15b后,成功表达并纯化了15kDa的Cq-FABP,并采用了多种荧光检测技术研究了不同脂肪源饲料中的主要脂肪酸与红螯光壳螯虾Cq-FABP的结合特性。利用ANS作为荧光标记,分别通过稳态荧光发射光谱、荧光共振能量转移(FRET)和时间分辨荧光研究了脂肪酸与Cq-FABP的结合。结果显示ANS除了能结合在Cq-FABP的脂肪酸结合位点外,还能结合在蛋白的其他位置。稳态荧光光谱结果显示Cq-FABP与棕榈酸(PA)、油酸(OA)、亚油酸(LA)、二十二碳五烯酸(EPA)和二十二碳六烯酸(DHA)的平衡解离常数Kd分别为1.1、2.4、3.1、2.9和5.8μM,Trp与ANS间的FRET效率约为10%；时间分辨荧光结果显示5μMCq-FABP在Buffer B中色氨酸残基的平均荧光寿命(τm)值约为3.04 ns,在ANS200μM时τm值缩短至2.00 ns,在加入足量的PA替代脂肪酸结合位点上的ANS后τm值恢复至2.33 ns,时间分辨荧光测得的FRET效率也在10%左右,与稳态荧光FRET结果相同。采用BODIPY C16与多种脂肪酸进行竞争性滴定,通过稳态荧光异向性检测获得的PA、OA、LA、EPA和DHA Kd值分别为1.8、2.7、3.8、3.7和7.2μM。结果显示Cq-FABP与脂肪酸亲和力与脂肪酸碳链长度和构型相关,碳链越长亲和力越差,弯曲型的脂肪酸的亲和力较高。Cq-FABP与脂肪酸的亲和力差异进一步证实了我们之前的推测：豆油组较高的Vg表达量可能是由于亚油酸通过控制FABP表达量的变化间接参与了基因表达的调控。
Since its introduction in China in 1992, demand for the red claw crayfish (Cherax quadricarinatus), a popular commercial aquaculture crustacean species, has increased dramatically due to its large size and positive marketing potential. In China, red claws are often fed commercial prawn feeds which despite their high price fail to meet the nutritional demands of the red claw. Feed is often one of highest costs in an aquaculture enterprise, and though diets that promote red claw growth is of growing interest to producers, little attention has been placed on developing adequate diets for maturing broodstock. Thus, nutritionally tailored and economical feed formulations are needed for red claws in aquaculture. In current study, the effects of four dietary lipid sources on growth and gonad maturation of pre-adult red claws were evaluated first. The effects of different cholesterol levels in feed were evaluated by the same way. To investigate the modulation function of different fatty acids, the fatty acid binding protein Cq-FABP was cloned and expressed. Some bio-photonics approaches were employed to study the ligands binding properties of Cq-FABP.
1. Effect of different dietary lipid sources on growth and biochemical composition of pre-adult female Cherax quadricarinatus
The effects of four dietary lipid sources (fish oil, peanut oil, soybean oil, pork lard) on growth and biochemical composition of pre-adult female red claw crayfish, Cherax quadricarinatus, were evaluated. Performance was evaluated by weight gain, specific growth rate (SGR), survival, feed conversion ratio (FCR), gonadosomatic index (GSI) and fatty acid composition. The proportion ofsaturated fatty acids (SFA), mono-unsaturated fatty acids (MUFA), and poly-unsaturated fatty acids (PUFA) in hepatopancreas varied as a result of lipid source, while differences in muscle were limited to MUFA levels. The optimal growth observed in red claws receiving the soybean oil diet. Conversely, diets high in EPA (20:5n3) and DHA (22:6n3) produced suboptimal results, suggesting that the importance of maturation diets with high EPA and DHA content may be exaggerated. As the dietary lipid requirements of growing and reproductively active red claw crayfish were satisfied by a plant oil that contained high levels of 18-carbon unsaturated fatty acids, soybean oil may be an economic alternative to the expensive lipid-fortified feed currently utilized by the aquaculture industry.
2. Effect of different dietary lipid sources on digestive enzyme activities and Vg expression of pre-adult female Cherax quadricarinatus
The effects of four dietary lipid sources (fish oil, peanut oil, soybean oil, pork lard) on digestive enzyme activities and Vg expresson of pre-adult female red claw crayfish, Cherax quadricarinatus, were evaluated. The activities of digestive enzymes were influenced by different dietary lipid sources, differed significantly. The soybean oil group has a higher activities than those from the other treatments. The low activities in red claws receiving the diet 4 possibly reflecting a lack of essential fatty acids in pork lard. Since GSI increases slowly at the onset of ovarian maturation, differences in dietary lipid sources did not alter GSI in present study, while the change in the hepatopancreatic Vg expression is significant. The Vg expression, with peak expression observed in red claws receiving the soybean oil diet, indicated that the fatty acids in soybean oil may have the function on stimulating the gonad maturation.
3. Effect of different dietary cholesterol levels on growth and reproduction of female pre-adult Cherax quadricarinatus
The effect of dietary cholesterol level on growth and reproduction performance of female pre-adult red claw crayfish Cherax quadricarinatus was evaluated over a 60-day culture period. Five experimental diets, supplemented with 0%,0.25%,0.50%, 0.75% and 1.00% cholesterol were evaluated. The growth performance, digestive enzyme activities and Vg expression in hepatopancreas were measured. A quadratic equation was used for regression analysis which was used to determine the digestive enzyme activities response to the dietary cholesterol level. The results showed that the female red claw crayfish fed with the dietary cholesterol ranged from 0.25-0.50% had better weight gain, SGR, survival rate, and FCR, and higher Lipase and Trypsin activities than those from the other treatments. The regression results from the growth performance and the digestive enzyme activities showed that the optimum cholesterol content was 0.55%. Female red claw crayfish fed with the diet without cholesterol supplementation had the lowest growth, digestive enzyme activities and the Vg expression. Though the group 5 (1% cholesterol) had the highest Vg expression, consider the overall performance, the recommended dietary cholesterol level for optimum growth and gonad maturation for female red claw crayfish under these environmental conditions was 0.55%. During rapid gonadal development phase in female C. quadricarinatus, a 0.75% cholesterol diet was recommended.
4. Molecular cloning and sequence analysis of the fatty acid binding protein (Cq-FABP) gene in female red claw crayfish Cherax quadricarinatus
A cDNA encoding red claw crayfish fatty acid binding protein (Cq-FABP) gene was cloned by using rapid amplification of cDNA ends (RACE). The full length cDNA was 772 bp, containing a 82 bp 5'-UTR region, a 291bp 3'-UTR region and a 399bp ORF region which encoded a 132 aa polypeptide. The secondary and tertiary structure assay showed that the Cq-FABP has 10β-strands and two a-helices, which is delimiting a cavity where the hydrophobic ligands are bound just as other FABPs. The deduced amino acids sequence of Cq-FABP sharing 90.9% homology with P.leniusculus and 56.8%,56.8%,53.8%,55.3%,53.8% with E.sinensis, P.clarkii, M.ensis, L.vannamei, P.monodon respectively. The homology with FABPs of human were lower, sharing 37.0%,35.4%,35.4%,38.6%,37.0%,40.9%,31.5%,38.6% homology with A-FABP, S-FABP, E-FABP, H-FABP, B-FABP, L-FABP, M-FABP, respectively. The high homology of Cq-FABP with B-FABP implied that they may have a close function on modulation of gene transcription. A NJ poylogenetic tree was also constructed based on reported FABPs amino acid sequences, results showed that the tree topology is agreement with traditional taxonomic relationships. The Decapoda species were in the same cluster, but the Cq-FABP together with the FABP from P.leniusculus were clustered in a different group with Decapoda species.
5. Expression pattern of the fatty acid binding protein (Cq-FABP) gene in female red claw crayfish Cherax quadricarinatus
A real-time qRT-PCR was employed to investigate the distribution of Cq-FABP in different tissues as well as to assess expression in hepatopancreas fed diets with different lipid sources. The beta-actin was used to normalize the Cq-FABP expression. Results showed that Cq-FABP was widely distributed, with the highest expression level in hepatopancreas, small amount of expression in intestine, detectable expression level in ovary, antennal gland, hemolymph, gills, stomach and testis, while expression was almost undetectable in muscle. The distribution manner indicated the possible function of Cq-FABP mainly involved in the intake and utilization of fatty acids. The Cq-FABP expression pattern in hepatopancreas with different lipid source diets treatment showed the influence of fatty acids on the Cq-FABP expression levels, the expression level seems relate to the amount of the long chain fatty acids in the diets, except the soybean oil group. Considering the good growth and high Vg expression level, the different expression pattern of Cq-FABP in soybean oil group implied the Cq-FABP may involved in gene modulation.
6. Expression, purification and fatty acids binding properties of the recombinant Cq-FABP
The Fragment contained the complete coding sequence of the Cq-FABP with NdeⅠand BamHⅠwas obtained by PCR, and an expression vector pET15b-His6-Cq-FABP has been constructed. The 15 kDa Cq-FABP protein was successfully expressed by the constructed Cq-FABP expression strain after the IPTG induction.
Interaction of various fatty acids with recombinant Cq-FABP was studied by different fluorescence procedures. In order to characterize the binding of several non fluorescent fatty acids (the dominant fatty acid in different lipid source diets) to Cq-FABP, we first indirectly determined their apparent affinities by an equilibrium displacement approach using ANS by using the steady-state fluorescence emission spectra. After correction by both the inner filter effect and the "non specific" binding of ANS, the Kd of palmitic acid (PA) was calculated from linear regression plotted by a series of competitive titrations. Next, a fluorescence resonance energy transfer (FRET) assay was conducted to address the binding of palmitic acid (PA) to Cq-FABP. TheΔ%FRET values were measured both by the steady-state fluorescence measurements and time-resolved fluorescence measurements. By plotting both the fluorescence intensity parameter and the FRET efficiency parameter, the calculated Kd,PA values were compatible with the one determined by monitoring the ANS fluorescence. After comparing the R-square of the two procedures, the Kd values of other fatty acids were measured by the first one. Results showed that the Kd values for Palmitic acid (PA), Oleic acid (OA), linoleic acid (LA), Eicosapentaenoic acid (EPA) and cis-4,7,10,13,16,19-Docosahexaenoic acid (DHA) were 1.1,2.4,3.1,2.9 and 5.8 respectively. Then the BODIPY-labeled fatty acids BODIPY C16 which does not have the "non-specific" with Cq-FABP was employed to measure the Kd values by steady-state fluorescence anisotropy based assay. The the Kd values for PA, OA, LA, EPA and DHA were 1.8,2.7,3.8,3.7 and 7.2 respectively, which were compatible with the value obtained by the previous procedure. Based on the Kd values of fatty acids, the Cq-FABP expression profile in the treatments with different lipid sources indicated that the high Vg expression may due to the LA, which may have the function of modulating the gene expression by the manner of regulating the Cq-FABP.
1.不同脂肪源饲料对雌性红鳌光壳螯虾生长及组织生化组成的影响
取平均体重为19.25±0.72g雌性红螯光壳螯虾,以鱼油、花生油、豆油和猪油作为脂肪源配制4种粗蛋白含量为33%,粗脂肪为8.5%的饲料,并以商业饲料作为对照投喂60天,研究不同脂肪源条件对雌性红螯光壳螯虾亲虾生长及组织生化组成的影响。结果显示：不同脂肪源饲料对雌性红螯光壳螯虾亲虾的增重、特殊生长率、存活率、饲料系数均有显著影响,而在本实验中对GSI无显著影响。其中以豆油作为脂肪源的饲料投喂的亲虾在增重、存活率等指标上均显著优于其他各组。在基本生化组成上,除了花生油组亲虾在肌肉和肝胰腺中脂肪和蛋白含量上略低外,其余各组无显著差异。在脂肪酸组成上,肌肉中主要的脂肪酸为c16:0, c18:0, c18:1n9, c18:2n6 and c20:5n3,在肝胰腺中为c16：0,c18：ln9,c18：2n6,肌肉中的c20:5n-3 (EPA)含量显著高于肝胰腺。不同饲料组对肝胰腺中的脂肪酸组成影响非常显著,饲料中SFA/MUFA/PUFA的比例直接影响了肝胰腺中脂肪酸的组成。各组肌肉脂肪酸的SFA/PUFA占总脂的比率没有显著的差异,仅MUFA有一定差异,其中第5组的的MUFA含量在总脂中所占比例最小。结果显示,在粗蛋白含量为33%,粗脂肪为8.5%条件下采用豆油作为脂肪源较适于此阶段的雌性红螯光壳螯虾生长。
2.不同脂肪源饲料对雌性红鳌光壳螯虾消化酶活力以及Vg基因表达的影响
在以不同脂肪源饲料投喂60天后,通过对雌性红螯光壳螯虾亲虾肝胰腺中消化酶活力以及卵黄蛋白源Vg基因的表达量的检测,研究不同脂肪源饲料对雌性亲虾的影响。结果显示消化酶活力受到不同脂肪源的影响,其中鱼油组、豆油组和商业饲料组脂肪酶活力显著高于花生油组和猪油组,但三者间无显著性差异；胰蛋白酶则以鱼油组、花生油组和豆油组较高,猪油组和商业饲料组相对较底；不同脂肪源饲料对淀粉酶活力影响相对较小,除花生油组略高外,其余各组无显著性差异,在纤维素酶活力上,除猪油组显著较低外,其余各组无显著差异。总体上讲,豆油组各消化酶活力均较好。虽然不同脂肪源组亲虾在GSI上没有显著差异,在肝胰腺Vg表达量上则有显著差异。其中豆油组投喂的亲虾最高的Vg表达量暗示了豆油较好的促卵巢成熟效果。结合生长指、消化酶活力和Vg表达,结果显示在本实验条件下以豆油作为脂肪源较适于雌性亲虾生长及生殖。
3.不同胆固醇含量饲料对红鳌光壳螫虾雌虾生长及生殖的影响
取平均体重为19.53+1.03g雌性红螯光壳螯虾,以0%、0.25%、0.5%、0.75%、1%胆固醇添加量配置5种粗蛋白含量为34%,粗脂肪为8.5%的饲料投喂60天。通过对不同实验组雌性红螯光壳螯虾亲虾的基本生长指标、脂肪酸组成、消化酶活力以及Vg表达量的研究,通过非线性回归来寻找此阶段雌性红螯光壳螯虾的胆固醇最适添加量。结果显示不同胆固醇添加量对肌肉和肝胰腺脂肪酸组成影响不显著,但对其基本生长指标、消化酶活力以及Vg表达均有显著影响。其中未添加组在增重、存活率等生长指标,脂肪酶、胰蛋白酶活力以及Vg表达量上均比添加胆固醇组低。在添加胆固醇的组别中,0.25%及0.5%的胆固醇添加组亲虾在增重率、SGR、存活率等生长指标,脂肪酶、胰蛋白酶活力以及Vg表达量上均显示出较好的效果,而高添加量组(1%)虽然在Vg表达量上最高,但在其他指标上则显示出对生长的抑制作用。对增重、SGR、存活率、FCR以及脂肪酶和胰蛋白酶活力的回归分析结果显示,此阶段雌虾胆固醇最适添加量为0.55%。因此,在雌性红螯光壳螯虾卵黄发生的中后期,可将胆固醇添加量增至0.75%以满足其卵巢快速发育期需求。
4.红螯光壳螯虾Cq-FABP基因的克隆和序列特征分析
通过兼并引物PCR获得部分片段后,采用RACE技术成功的克隆了红螯光壳螯虾脂肪酸结合蛋白FABP基因的全长cDNA序列。该序列全长772bp,包含一个82bp的5’-UTR区域和一个291bp的3’-UTR区域,开放阅读框长399bp,编码132个氨基酸。通过生物信息学方法对其二级、三级结构预测结果显示该蛋白具有FABP典型的特征：10个β折叠和2个α螺旋,并通过β折叠和α螺旋形成能够容纳疏水配基的桶装结构。同源性比对结果显示红螯光壳螯虾脂肪酸结合蛋白Cq-FABP与通讯螯虾具有最高的同源性,为90.9%；与其他已知的甲壳类十足目种类的同源性则相对较低,和中华绒螯蟹、克氏原螯虾、刀额新对虾、凡纳滨对虾以及斑节对虾FABP的同源性分别为56.8%、56.8%、53.8%、55.3%和53.8%。而与人类多种FABP的比对结果显示Cq-FABP与人类脑型脂肪酸结合蛋白B-FABP具有最高的同源性,为40.9%,与心型和肌肉型FABP的同源性次之,为38.6%,这也暗示了其可能具有功能的多样性。系统进化结果也显示红螯光壳螯虾FABP虽然与其他甲壳类十足日FABP同属于一簇,但其与通讯螯虾同属与一个分支,其他十足目种类则属于另一分支。
5.红螯光壳螯虾Cq-FABP基因的表达特征分析
采用实时定量PCR技术研究了红螯光壳螯虾Cq-FABP基因在不同组织中的表达以及受不同脂肪源饲料影响后在肝胰腺中的表达量变化,均以β-actin作为内参。结果发现虽然在多个组织中均检测到Cq-FABP表达,但主要在肝胰腺组织中表达。在肠中也有少量表达,但表达量仅为肝胰腺腺中的1/70,在卵巢、触角腺、血淋巴、腮、胃和精巢中也有极微量的表达,但在肌肉中几乎无表达。由于其表达量最高的在肝胰腺组织,其次在肠,暗示了Cq-FABP的功能可能主要涉及脂肪酸的摄取和利用。以不同脂肪源饲料投喂的亲虾肝胰腺中Cq-FABP的表达量与脂肪源中的长链脂肪酸比例呈正比,但长链脂肪酸比例较为接近的花生油和豆油组表达量却较低。根据豆油组的亲虾的生长情况及的Vg高表达量推测可能与Cq-FABP的表达有关,而Cq-FABP与人B-FABP的最高同源性也暗示了不同的脂肪酸可能可以通过调节Cq-FABP的方式参与到信号调控中。
6. Cq-FABP蛋白的表达纯化及与脂肪酸结合特性的研究
将红螯光壳螯虾脂肪酸结合蛋白Cq-FABP基因重组入原核表达载体pET15b后,成功表达并纯化了15kDa的Cq-FABP,并采用了多种荧光检测技术研究了不同脂肪源饲料中的主要脂肪酸与红螯光壳螯虾Cq-FABP的结合特性。利用ANS作为荧光标记,分别通过稳态荧光发射光谱、荧光共振能量转移(FRET)和时间分辨荧光研究了脂肪酸与Cq-FABP的结合。结果显示ANS除了能结合在Cq-FABP的脂肪酸结合位点外,还能结合在蛋白的其他位置。稳态荧光光谱结果显示Cq-FABP与棕榈酸(PA)、油酸(OA)、亚油酸(LA)、二十二碳五烯酸(EPA)和二十二碳六烯酸(DHA)的平衡解离常数Kd分别为1.1、2.4、3.1、2.9和5.8μM,Trp与ANS间的FRET效率约为10%；时间分辨荧光结果显示5μMCq-FABP在Buffer B中色氨酸残基的平均荧光寿命(τm)值约为3.04 ns,在ANS200μM时τm值缩短至2.00 ns,在加入足量的PA替代脂肪酸结合位点上的ANS后τm值恢复至2.33 ns,时间分辨荧光测得的FRET效率也在10%左右,与稳态荧光FRET结果相同。采用BODIPY C16与多种脂肪酸进行竞争性滴定,通过稳态荧光异向性检测获得的PA、OA、LA、EPA和DHA Kd值分别为1.8、2.7、3.8、3.7和7.2μM。结果显示Cq-FABP与脂肪酸亲和力与脂肪酸碳链长度和构型相关,碳链越长亲和力越差,弯曲型的脂肪酸的亲和力较高。Cq-FABP与脂肪酸的亲和力差异进一步证实了我们之前的推测：豆油组较高的Vg表达量可能是由于亚油酸通过控制FABP表达量的变化间接参与了基因表达的调控。
Since its introduction in China in 1992, demand for the red claw crayfish (Cherax quadricarinatus), a popular commercial aquaculture crustacean species, has increased dramatically due to its large size and positive marketing potential. In China, red claws are often fed commercial prawn feeds which despite their high price fail to meet the nutritional demands of the red claw. Feed is often one of highest costs in an aquaculture enterprise, and though diets that promote red claw growth is of growing interest to producers, little attention has been placed on developing adequate diets for maturing broodstock. Thus, nutritionally tailored and economical feed formulations are needed for red claws in aquaculture. In current study, the effects of four dietary lipid sources on growth and gonad maturation of pre-adult red claws were evaluated first. The effects of different cholesterol levels in feed were evaluated by the same way. To investigate the modulation function of different fatty acids, the fatty acid binding protein Cq-FABP was cloned and expressed. Some bio-photonics approaches were employed to study the ligands binding properties of Cq-FABP.
1. Effect of different dietary lipid sources on growth and biochemical composition of pre-adult female Cherax quadricarinatus
The effects of four dietary lipid sources (fish oil, peanut oil, soybean oil, pork lard) on growth and biochemical composition of pre-adult female red claw crayfish, Cherax quadricarinatus, were evaluated. Performance was evaluated by weight gain, specific growth rate (SGR), survival, feed conversion ratio (FCR), gonadosomatic index (GSI) and fatty acid composition. The proportion ofsaturated fatty acids (SFA), mono-unsaturated fatty acids (MUFA), and poly-unsaturated fatty acids (PUFA) in hepatopancreas varied as a result of lipid source, while differences in muscle were limited to MUFA levels. The optimal growth observed in red claws receiving the soybean oil diet. Conversely, diets high in EPA (20:5n3) and DHA (22:6n3) produced suboptimal results, suggesting that the importance of maturation diets with high EPA and DHA content may be exaggerated. As the dietary lipid requirements of growing and reproductively active red claw crayfish were satisfied by a plant oil that contained high levels of 18-carbon unsaturated fatty acids, soybean oil may be an economic alternative to the expensive lipid-fortified feed currently utilized by the aquaculture industry.
2. Effect of different dietary lipid sources on digestive enzyme activities and Vg expression of pre-adult female Cherax quadricarinatus
The effects of four dietary lipid sources (fish oil, peanut oil, soybean oil, pork lard) on digestive enzyme activities and Vg expresson of pre-adult female red claw crayfish, Cherax quadricarinatus, were evaluated. The activities of digestive enzymes were influenced by different dietary lipid sources, differed significantly. The soybean oil group has a higher activities than those from the other treatments. The low activities in red claws receiving the diet 4 possibly reflecting a lack of essential fatty acids in pork lard. Since GSI increases slowly at the onset of ovarian maturation, differences in dietary lipid sources did not alter GSI in present study, while the change in the hepatopancreatic Vg expression is significant. The Vg expression, with peak expression observed in red claws receiving the soybean oil diet, indicated that the fatty acids in soybean oil may have the function on stimulating the gonad maturation.
3. Effect of different dietary cholesterol levels on growth and reproduction of female pre-adult Cherax quadricarinatus
The effect of dietary cholesterol level on growth and reproduction performance of female pre-adult red claw crayfish Cherax quadricarinatus was evaluated over a 60-day culture period. Five experimental diets, supplemented with 0%,0.25%,0.50%, 0.75% and 1.00% cholesterol were evaluated. The growth performance, digestive enzyme activities and Vg expression in hepatopancreas were measured. A quadratic equation was used for regression analysis which was used to determine the digestive enzyme activities response to the dietary cholesterol level. The results showed that the female red claw crayfish fed with the dietary cholesterol ranged from 0.25-0.50% had better weight gain, SGR, survival rate, and FCR, and higher Lipase and Trypsin activities than those from the other treatments. The regression results from the growth performance and the digestive enzyme activities showed that the optimum cholesterol content was 0.55%. Female red claw crayfish fed with the diet without cholesterol supplementation had the lowest growth, digestive enzyme activities and the Vg expression. Though the group 5 (1% cholesterol) had the highest Vg expression, consider the overall performance, the recommended dietary cholesterol level for optimum growth and gonad maturation for female red claw crayfish under these environmental conditions was 0.55%. During rapid gonadal development phase in female C. quadricarinatus, a 0.75% cholesterol diet was recommended.
4. Molecular cloning and sequence analysis of the fatty acid binding protein (Cq-FABP) gene in female red claw crayfish Cherax quadricarinatus
A cDNA encoding red claw crayfish fatty acid binding protein (Cq-FABP) gene was cloned by using rapid amplification of cDNA ends (RACE). The full length cDNA was 772 bp, containing a 82 bp 5'-UTR region, a 291bp 3'-UTR region and a 399bp ORF region which encoded a 132 aa polypeptide. The secondary and tertiary structure assay showed that the Cq-FABP has 10β-strands and two a-helices, which is delimiting a cavity where the hydrophobic ligands are bound just as other FABPs. The deduced amino acids sequence of Cq-FABP sharing 90.9% homology with P.leniusculus and 56.8%,56.8%,53.8%,55.3%,53.8% with E.sinensis, P.clarkii, M.ensis, L.vannamei, P.monodon respectively. The homology with FABPs of human were lower, sharing 37.0%,35.4%,35.4%,38.6%,37.0%,40.9%,31.5%,38.6% homology with A-FABP, S-FABP, E-FABP, H-FABP, B-FABP, L-FABP, M-FABP, respectively. The high homology of Cq-FABP with B-FABP implied that they may have a close function on modulation of gene transcription. A NJ poylogenetic tree was also constructed based on reported FABPs amino acid sequences, results showed that the tree topology is agreement with traditional taxonomic relationships. The Decapoda species were in the same cluster, but the Cq-FABP together with the FABP from P.leniusculus were clustered in a different group with Decapoda species.
5. Expression pattern of the fatty acid binding protein (Cq-FABP) gene in female red claw crayfish Cherax quadricarinatus
A real-time qRT-PCR was employed to investigate the distribution of Cq-FABP in different tissues as well as to assess expression in hepatopancreas fed diets with different lipid sources. The beta-actin was used to normalize the Cq-FABP expression. Results showed that Cq-FABP was widely distributed, with the highest expression level in hepatopancreas, small amount of expression in intestine, detectable expression level in ovary, antennal gland, hemolymph, gills, stomach and testis, while expression was almost undetectable in muscle. The distribution manner indicated the possible function of Cq-FABP mainly involved in the intake and utilization of fatty acids. The Cq-FABP expression pattern in hepatopancreas with different lipid source diets treatment showed the influence of fatty acids on the Cq-FABP expression levels, the expression level seems relate to the amount of the long chain fatty acids in the diets, except the soybean oil group. Considering the good growth and high Vg expression level, the different expression pattern of Cq-FABP in soybean oil group implied the Cq-FABP may involved in gene modulation.
6. Expression, purification and fatty acids binding properties of the recombinant Cq-FABP
The Fragment contained the complete coding sequence of the Cq-FABP with NdeⅠand BamHⅠwas obtained by PCR, and an expression vector pET15b-His6-Cq-FABP has been constructed. The 15 kDa Cq-FABP protein was successfully expressed by the constructed Cq-FABP expression strain after the IPTG induction.
Interaction of various fatty acids with recombinant Cq-FABP was studied by different fluorescence procedures. In order to characterize the binding of several non fluorescent fatty acids (the dominant fatty acid in different lipid source diets) to Cq-FABP, we first indirectly determined their apparent affinities by an equilibrium displacement approach using ANS by using the steady-state fluorescence emission spectra. After correction by both the inner filter effect and the "non specific" binding of ANS, the Kd of palmitic acid (PA) was calculated from linear regression plotted by a series of competitive titrations. Next, a fluorescence resonance energy transfer (FRET) assay was conducted to address the binding of palmitic acid (PA) to Cq-FABP. TheΔ%FRET values were measured both by the steady-state fluorescence measurements and time-resolved fluorescence measurements. By plotting both the fluorescence intensity parameter and the FRET efficiency parameter, the calculated Kd,PA values were compatible with the one determined by monitoring the ANS fluorescence. After comparing the R-square of the two procedures, the Kd values of other fatty acids were measured by the first one. Results showed that the Kd values for Palmitic acid (PA), Oleic acid (OA), linoleic acid (LA), Eicosapentaenoic acid (EPA) and cis-4,7,10,13,16,19-Docosahexaenoic acid (DHA) were 1.1,2.4,3.1,2.9 and 5.8 respectively. Then the BODIPY-labeled fatty acids BODIPY C16 which does not have the "non-specific" with Cq-FABP was employed to measure the Kd values by steady-state fluorescence anisotropy based assay. The the Kd values for PA, OA, LA, EPA and DHA were 1.8,2.7,3.8,3.7 and 7.2 respectively, which were compatible with the value obtained by the previous procedure. Based on the Kd values of fatty acids, the Cq-FABP expression profile in the treatments with different lipid sources indicated that the high Vg expression may due to the LA, which may have the function of modulating the gene expression by the manner of regulating the Cq-FABP.
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