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褪黑激素影响山羊绒生长的分子机理
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摘要
通过对内蒙古绒山羊埋植褪黑激素(MT)试验,研究连续2年埋植MT、只埋植1年和停止埋植对其绒毛生产性能和皮肤中有关绒毛生长相关基因表达量变化的影响,进而为研究绒毛生长的分子机理提供可参考的依据,也为MT在生产上的运用奠定理论基础。为研究MT对绒毛生长特性与绒毛品质的影响,每月测定受试羊绒和毛细度、长度及绒毛质量比;同时结合荧光定量PCR实验和原位杂交实验,研究埋植MT对绒毛生长相关基因在绒山羊皮肤毛囊中表达的影响。研究结果发现:
     埋植MT缩短绒毛生长周期。从7月份开始埋植MT经过1年的埋植出现1年内2次脱绒现象,而再持续埋植1年MT出现2年内4次脱绒现象,然而埋植1年MT后7月份停止埋植将在2年内出现3次脱绒现象,并推迟其翌年春季脱绒时间。虽然秋季脱落的绒比春季抓绒时的绒短,但其绒长度也能满足生产需要。埋植MT或停止埋植对成年羊春季抓绒时绒厚度、长度和细度无影响;对羔羊连续2年埋植MT,使其抓绒时绒细度变细,而埋植1年后停止埋植MT具有绒长度变短的趋势。埋植MT对粗毛长度和生长周期无影响,但对周岁羊埋植MT的当年其粗毛有变细的趋势。从羔羊阶段开始连续2年埋植MT明显降低周岁羊抓绒时的绒毛比,但埋植MT具有提高成年羊抓绒时的绒毛比的趋势。开始埋植MT的当年出现产绒量提高的趋势,而连续2年埋植MT后导致第二年产绒量比第一年的产绒量下降;停止埋植MT后,其产绒量变化趋势与对照组类似。
     通过RT-PCR实验研究发现:在内蒙古成年绒山羊皮肤组织中,毛囊生长前期提高表达量的基因:TGFβ2、Deltal、Notch1、Wnt10b、MSX-2、β-catenin、 FGF5、PDGFA、PDGFRa和RORa。绒毛快速生长期低表达的基因:TGFβ2、TGFPRⅡ、Notch1、hairless、WntlOb、β-catenin、PDGFA、NTRK3、RORa、RORβ和RORγ。退行期至休止期初提高表达量的基因:BMPR1B、TGFβ2、Notch1、hairless、 Wnt10b、β-catenin、PDGFA、NTRK3、RORα、RORβ和RORy。退行期提高后休止期初降低表达量的基因:BMP2和Deltal基因。休止期高表达的基因:BMP2、TGFβRⅡ、PDGFA、NTRK3、RORα、 RORβ和RORy基因。
     停止埋植MT组和连续2年埋植MT组在绒毛快速生长期和休止初期Wnt10b基因表达量的下降,可能与埋植MT促使毛囊进入休止期有关。然而连续埋植1年或2年MT在休止期后期提高Wnt10b基因的表达量、降低BMP2基因的表达量,从而诱导毛囊提前进入了新一轮的生长期。埋植MT可能通过影响RORy基因的表达来影响Wnt10b的表达,从而降低皮肤中Wnt10b与TGFβ2基因表达量的相关性,从而影响了其它信号通路分子的表达。与对照组相比MT处理组降低毛囊退行期和休止期RORα、RORβ和RORy基因,休止初期β-catenin基因,以及NTRK3基因休止期的表达量;提高休止初期MSX-2和FGF5基因的表达量。其中,停止埋植MT后也降低绒毛快速生长期TGFβRⅡ、NTRK3、Deltal和RORa基因的表达量,提高休止后期TGFβRⅡ和BMPRIB基因的表达量。连续2年埋植MT也降低快速生长期BMP2基因,休止期TGFβRⅡ基因的表达量;提高BMPR1B、Notch1和hairless基因在休止前期的表达量。对成年羊7月份开始埋植MT也降低退行期BMP2基因和休止期TGFβRⅡ基因的表达量。Wnt10b、PDGFA和RORa基因为绒毛生长周期相关基因,埋植MT影响毛囊生长周期,而这些基因在皮肤组织中的表达模式也随着毛囊周期而变化。在皮肤毛囊组织中,TGFβ信号通路通过WNT信号通路调节其它信号通路,其中TGFβ信号通路中的BMP2和TGFβ2基因在整个毛囊周期性生长过程中具有很重要的作用。
     原位杂交研究发现,初级毛囊和次级毛囊上BMP2基因mRNA的表达一样;埋植MT使其全年表达在外根鞘和连接组织鞘上。脱绒后,埋植MT使Wnt10b基因mRNA提前1个月在次级毛囊外根鞘上表达,提前2个月在其连接组织鞘上表达。
The paper studies the implanting melatonin (MT) of Cashmere goats in Inner Mongolia. The changes in production characters and the expression level of growth related gene in skin were studied for three treatments:implanted MT for two consecutive years, implanting MT for1year and stopped implant MT. The analysis results could provide references and basis for the molecular mechanism of Cashmere growth research, and establish the theoretical foundation for the actual production practice of MT. To study the effect of MT on the Cashmere growth traits and the quality of Cashmere, diameter and length of Cashmere and wool were measured monthly; at the same time, study on the influence of implanting MT on the expression level of growth related gene in skin using the RT-PCR and in-situ hybridization techniques. Those results suggested that:
     The implanting MT shortened the time of Cashmere growth cycle. Starting to implant MT from July, the Cashmere goats depilated Cashmere twice in one year, and four times during two years after implanting MT another year. Stopped implanting MT after the first one year, the Cashmere goats depilated Cashmere three times during two years and delayed depilation in next spring. Although the depilated Cashmere in autumn was shorter than in spring, the length of Cashmere can meet the requirement of production. Implanting MT or stopped implanting had no effect on Cashmere thickness, length and diameter of mature goats in spring; implanting MT for two consecutive years to lamb can decrease the Cashmere diameter. Stopped implanting after one year treatment for lamb made the Cashmere length getting shorter. Implanting MT had no effect on wool length and growth cycle, but decreased the tendency of wool diameter for one year old Cashmere goat in implanting year. The proportion of cashmere and wool had been significantly declined by consecutive implanting MT for two years in lamb, had been improved by implanting MT in mature goat. The Cashmere field improved in the first year of implanting MT, and decreased in the next year after implanting MT for two consecutive years; the Cashmere field had similar trend to control group after stopping implanting MT for Cashmere goat.
     RT-PCR results showed:In the mature goat skin, the expression levels of TGFβ2, Deltal, Notch1, Wnt10b, MSX-2, p-catenin, FGF5, PDGFA, PDGFRa and RORa had increased during Propagate anagen; TGFβ2, TGFβRⅡ, Notch1, hairless, Wnt10b, P-catenin, PDGFA, NTRK3, RORa, RORPβ和RORγ had decreased on fast growing stage of Cashmeres; BMPR1B, TGFβ2, Notch1, hairless, Wnt10b, β-catenin, PDGFA, NTRK3, RORα, RORβ and RORγ had decreased in catagen and refractory telogen; BMP2and Delta1had decreased in refractory telogen after increased in telegon; BMP2, TGFβRⅡ, PDGFA:NTRK3, RORa, RORβ and RORγ had highest in categon.
     It may related to promote hair follicle cycle in the categon, that Wnt10b gene expression level in mature goat skin was decreased after stopped implanting MT or consecutive implanting MT for two years in Cashmere fast growing stage and refractory telogen. However, implanting MT for1or2years increased the expression level of Wnt10b and decreased the expression level of BMP2in telogen, which induced hair follicle growth initiate earlier. Implanting MT influence the expression of Wnt10b by affecting the expression of RORy gene, this decreased the relativity between Wnt10b and RORy gene, further influence the expression of other signal pathway molecular. In contrast with control group, implanting MT decrease the expression level of RORa, RORP and RORy gene in catagen and telogen, β-catenin gene in refractory telogen and NTRK3gene in telogen, however increased the expression level of MSX-2and FGF5gene. The expression levels of TGFβRⅡ, NTRK3, Delta1and RORa gene decreased in hair fast growing stage by stopped implanting MT, but the expression level of TGFβRⅡ and BMPR1B gene in competent telogen was decreased. For the consecutive implanting MT group, the expression level of BMP2gene in anagen and the expression level of TGFβRⅡ gene in telogen were decreased; however increased the expression level of BMPR1B, Notch1and hairless gene in refractory telogen. The expression level of BMP2gene in cetagon and the expression level of TGFβRⅡ gene in telogon were decreased by implanting MT from july to mature goat. Wnt10b, PDGFA and RORa gene were related to hair follicle growth cycle which was influenced by implanting MT. The expression level of WntlOb, PDGFA and RORa gene were changed with hair follicle cycle. In the skin, the other signal pathways were regulated by TGFβ signal pathway which through WNT signals.
     The results of In Situ Hybridization indicated that there was the same expression pattern for BMP2mRNA on the two kind of hair follicle; While BMP2was expressed on Outer Root Sheath and Connective Tissue Sheath over year after implanting MT. Wnt10b mRNA was detected one month earlier on Outer Root Sheath and two months earlier on Connective Tissue Sheath for Secondary hair follicle after fleecing for implanting group than control group.
引文
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