Per2和c-fos mRNA及FOS蛋白在缰核内的节律性表达
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摘要
下丘脑视交叉上核(SCN)和松果体被认为是调控昼夜节律的两个重要结构,丘脑的缰核(Hb)密切联系这两个部位,并且Hb可直接接受视网膜纤维的投射,Hb神经元的放电活动在离体条件下具有昼夜节律性。本课题分别以大鼠全缰核(Hb)及内侧缰核(MHb)、外侧缰核(LHb)为研究对象,在有光照信号(LD)及缺乏光照信号(DD)条件下,使用半定量RT-PCR方法及免疫组织化学方法,检测了正常及SCN损毁的大鼠全缰核每昼夜两个时间点(13:00时和1:00时)和内、外侧缰核每昼夜六个时间点(13:00时,17:00时,21:00时,1:00时,5:00时,9:00时)昼夜节律基因Per2和即刻早期基因c-fos及其蛋白的表达。
本实验结果显示:在LD条件下,Per2和c-fos基因及FOS蛋白的表达在全缰核和LHb内显示出昼高夜低的节律振荡,在MHb内无明显的节律性;当SCN损毁后,Per2和c-fos基因及FOS蛋白的表达无论是全缰核还是LHb和MHb内均没有节律性。
在DD条件下,取消了外界光照对视网膜的刺激后,Per2和c-fos基因及FOS蛋白的表达在全缰核、LHb和MHb内均显示出昼低夜高的振荡,表现出与LD光照条件下完全相反的变化;当SCN损毁后,尽管在LHb内有些时间点的表达差异消失,Per2和c-fos基因及FOS蛋白的表达仍然维持其在全缰核和LHb内昼低夜高节律性的表达,而Per2和c-fos基因及FOS蛋白的表达在MHb内的节律性振荡消失。
综上所述,Hb自身的节律直接/间接受控于SCN,它可能做为SCN的次级位点参与中枢昼夜节律的调控,是中枢昼夜节律控制系统中的重要成分。
Circadian rhythm is the rhythm with 24h periodic variations which is a basic vital phenomena. Disruption of circadian rhythm will lead to functional disorder even disease. The suprachiasmatic nucleus of the hypothalamus (SCN) is the master circadian pacemaker or clock in the mammalian brain. A key feature of a circadian pacemaker is its ability to entrain to changing environmental cues. The principal entraining stimulus in mammals is the LD cycle, with photic information being conveyed to the SCN via the direct pathway-glutamatergic retinohypothalamic tract (RHT) or other indirect pathways to coordinate the whole circadian system and control the most activities of the bodies such as physiological, metabolic and behavioural. Habenular nucleus(Hb)which locates epithalamus is functionally interposed between limbic and striatal systems through the involvement of the midbrain structures. Hb is the cross point of cardiovascular activity, sleep-wake cycle, production, body temperature and behabior which all show daily rhythmicity. Our previous study showed that the discharge of habenula neurons had obvious circadian rhythm. Cells in the Hb had higher baseline firing rates during the day than during the night in vivo and vitro, while photic responses were of significantly larger amplitude among LHb cells duringthe projected night than during the projected day. Mahboubeh found the Number of c-FOS were higher during the subjective night-time than that during the subjectve day-time in Hamsters LHbM in constant darkness (DD). So the habenula may be one important component in linking circadian and motivational systems and may contribute to the regulation of these systems, as well as to the rhythmicity of their function. Yet it is not reported about the molecular mechanism of daily rhythmicity in the Hb till now. So many experiments found that the subtypes of Period(Per) family: Period1(Per1) and Period2(Per2) play the fundamental effects in creating circadian rhythm. Particularly Per2 plalys more important roles.
Methods:
Hb was removed on 13: 00 and 1: 00 and LHb and MHb were separated on 13: 00, 17: 00, 21: 00, 1: 00, 5: 00 and 9: 00 from normal rats and SCN lesioned rats. Immunohistochemical staining was used to detect the expression of FOS protein and Per2 and c-fos mRNA expression was determined with RT-PCR method on the time point as shown above.
Results:
1. The expression of Per2 and c-fos mRNA and FOS protein in the Hb:
1)The expression of Per2 and c-fos mRNA and FOS protein in the Hb showed the rhyhmicity with higher expression during the projective day than night in LD cycle(P<0.05). And this rhythmicity will disappear when SCN was lesioned(P>0.05)。
2)The expression of Per2 and c-fos mRNA and FOS protein in the Hb showed the rhyhmicity with higher expression during the subjective night than the subjective day in DD cycle when the photic stimulus from the environment was abolished(P<0.05), which was antipodal to the change in LD cycle. When SCN was lesioned, this rhythmicity could be maintained in the Hb(P<0.05)。
2. The expression of Per2 and c-fos mRNA and FOS protein in the LHb and MHb:
1)The expression of Per2 and c-fos mRNA and FOS protein in the LHb showed the rhyhmicity with higher expression during the projective day than night in LD cycle as in the Hb:The expression of Per2 mRNA was higher on 9: 00 and 13: 00 than that on 1: 00(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 13: 00 and 17: 00 than that on 1: 00(P<0.05); The expression of Per2 and c-fos mRNA and FOS protein in the MHb had no rhythmicity(P>0.05). The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned(P>0.05).
2)The expression of Per2 and c-fos mRNA and FOS protein in the LHb and MHb showed the rhyhmicity with higher expression during the subjective night than day in DD cycle when the photic stimulus from the environment was abolished as in the Hb: The expression of Per2 mRNA was lower on 9: 00 and 13: 00 than that on 1: 00(P<0.05)in the LHb(P<0.05) and the expression was also lower on 13: 00 than that on 21: 00(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 1: 00 than that on any time point during the whole 24h in the LHb(P<0.05); The expression of Per2 was higher on 1: 00 than that on any time point during the whole 24h in the MHb(P<0.05); and c-fos mRNA and FOS protein in the MHb had no rhythmicity(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 1: 00 than that on 13: 00 and 17: 00 in the MHb(P<0.05).
The expression of Per2 and c-fos mRNA and FOS protein in the LHb could be maintained the rhyhmicity with higher expression during the subjective night than day in DD cycle after SCN was lesioned: The expression was higher om 21: 00 and 1: 00 than that on 1: 00(P<0.05). The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned in the MHb(P>0.05).
Discussion:
The expression of Per2 and c-fos mRNA and FOS protein in the LHb and Hb showed the rhyhmicity with higher expression during the projective day than night, while there was no obvious rhythnicity in the MHb on different tine point in LD cycle. This rhythmic oscillation was coincident with the results from Prof. Zhao Hua who showed that cells in the LHb showed higher baseline firing rates during the day than during the night and cells in the MHb showed no rhythmic oscillation in vitro slice preparation. And in Zhao’s experiment of cells recorded in the LHb, 42% were tonically activated or suppressed by retinal illumination, while significantly fewer cells recorded in the MHb responded to retinal illumination(19%). It was reported that photosignal from the environment could project to the Hb directly and indirectly which made Hb receive the effect of photosignal. The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned in the LHb and Hb and there was still no rhythmicity in the MHb. Therefore, we can conclude that SCN is very significant to the Hb to maintain its rhythmicity in LD cycle; And because the rhythmic expression of Per2 and c-fos mRNA and FOS protein oscillate in the LHb coincident to the Hb, the rhythmicity of the LHb master that of the whole Hb.
The expression of Per2 and c-fos mRNA and FOS protein showed the rhyhmicity with lower expression during the subjective day than the subjective night no matter in the LHb, MHb or in the Hb on different time point in DD cycle. Of all our results, there was no significance between 17: 00 and 21: 00(2 h before lights off and after lights off)about the expression of FOS protein which was identical with the story from Mahboubeh. Although Mahboubeh also found that the FOS protein expressed low on the subjective day and high on the subjective night in LHbM of hamster in DD cycle, the mechanism is excluded. The expression of Per2 and c-fos mRNA and FOS protein in the LHb could be maintained the rhyhmicity with higher expression during the subjective night than the subjective day in DD cycle after SCN was lesioned. Although the expressive difference abolished among some time points, the difference between 13: 00 and 1: 00 still existed in the LHb; The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost in the MHb after SCN was lesioned. Thus, we can conclude that SCN is not indispensable to the Hb in DD; And the results futher support the phenomena: the rhythmicity of the LHb master that of the whole Hb.
Conclusion:
On the whole, there is a rhythmic oscillation in the LHb, MHb and Hb. Hb must rely on SCN in LD cycle, and SCN is very important. Although the expressive difference abolished among some time points in DD cycle, the expression of Per2 and c-fos mRNA and FOS protein still maintain the rhyhmicity with lower expression during day than the subjective night in the LHb, MHb or in the Hb. Hb can oscillates independent of SCN in DD cycle, but it is not a fully automic oscillation. Hb may participate the regulation of circadian rhythm system on the body and may play a different role from SCN in this system.
本实验结果显示:在LD条件下,Per2和c-fos基因及FOS蛋白的表达在全缰核和LHb内显示出昼高夜低的节律振荡,在MHb内无明显的节律性;当SCN损毁后,Per2和c-fos基因及FOS蛋白的表达无论是全缰核还是LHb和MHb内均没有节律性。
在DD条件下,取消了外界光照对视网膜的刺激后,Per2和c-fos基因及FOS蛋白的表达在全缰核、LHb和MHb内均显示出昼低夜高的振荡,表现出与LD光照条件下完全相反的变化;当SCN损毁后,尽管在LHb内有些时间点的表达差异消失,Per2和c-fos基因及FOS蛋白的表达仍然维持其在全缰核和LHb内昼低夜高节律性的表达,而Per2和c-fos基因及FOS蛋白的表达在MHb内的节律性振荡消失。
综上所述,Hb自身的节律直接/间接受控于SCN,它可能做为SCN的次级位点参与中枢昼夜节律的调控,是中枢昼夜节律控制系统中的重要成分。
Circadian rhythm is the rhythm with 24h periodic variations which is a basic vital phenomena. Disruption of circadian rhythm will lead to functional disorder even disease. The suprachiasmatic nucleus of the hypothalamus (SCN) is the master circadian pacemaker or clock in the mammalian brain. A key feature of a circadian pacemaker is its ability to entrain to changing environmental cues. The principal entraining stimulus in mammals is the LD cycle, with photic information being conveyed to the SCN via the direct pathway-glutamatergic retinohypothalamic tract (RHT) or other indirect pathways to coordinate the whole circadian system and control the most activities of the bodies such as physiological, metabolic and behavioural. Habenular nucleus(Hb)which locates epithalamus is functionally interposed between limbic and striatal systems through the involvement of the midbrain structures. Hb is the cross point of cardiovascular activity, sleep-wake cycle, production, body temperature and behabior which all show daily rhythmicity. Our previous study showed that the discharge of habenula neurons had obvious circadian rhythm. Cells in the Hb had higher baseline firing rates during the day than during the night in vivo and vitro, while photic responses were of significantly larger amplitude among LHb cells duringthe projected night than during the projected day. Mahboubeh found the Number of c-FOS were higher during the subjective night-time than that during the subjectve day-time in Hamsters LHbM in constant darkness (DD). So the habenula may be one important component in linking circadian and motivational systems and may contribute to the regulation of these systems, as well as to the rhythmicity of their function. Yet it is not reported about the molecular mechanism of daily rhythmicity in the Hb till now. So many experiments found that the subtypes of Period(Per) family: Period1(Per1) and Period2(Per2) play the fundamental effects in creating circadian rhythm. Particularly Per2 plalys more important roles.
Methods:
Hb was removed on 13: 00 and 1: 00 and LHb and MHb were separated on 13: 00, 17: 00, 21: 00, 1: 00, 5: 00 and 9: 00 from normal rats and SCN lesioned rats. Immunohistochemical staining was used to detect the expression of FOS protein and Per2 and c-fos mRNA expression was determined with RT-PCR method on the time point as shown above.
Results:
1. The expression of Per2 and c-fos mRNA and FOS protein in the Hb:
1)The expression of Per2 and c-fos mRNA and FOS protein in the Hb showed the rhyhmicity with higher expression during the projective day than night in LD cycle(P<0.05). And this rhythmicity will disappear when SCN was lesioned(P>0.05)。
2)The expression of Per2 and c-fos mRNA and FOS protein in the Hb showed the rhyhmicity with higher expression during the subjective night than the subjective day in DD cycle when the photic stimulus from the environment was abolished(P<0.05), which was antipodal to the change in LD cycle. When SCN was lesioned, this rhythmicity could be maintained in the Hb(P<0.05)。
2. The expression of Per2 and c-fos mRNA and FOS protein in the LHb and MHb:
1)The expression of Per2 and c-fos mRNA and FOS protein in the LHb showed the rhyhmicity with higher expression during the projective day than night in LD cycle as in the Hb:The expression of Per2 mRNA was higher on 9: 00 and 13: 00 than that on 1: 00(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 13: 00 and 17: 00 than that on 1: 00(P<0.05); The expression of Per2 and c-fos mRNA and FOS protein in the MHb had no rhythmicity(P>0.05). The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned(P>0.05).
2)The expression of Per2 and c-fos mRNA and FOS protein in the LHb and MHb showed the rhyhmicity with higher expression during the subjective night than day in DD cycle when the photic stimulus from the environment was abolished as in the Hb: The expression of Per2 mRNA was lower on 9: 00 and 13: 00 than that on 1: 00(P<0.05)in the LHb(P<0.05) and the expression was also lower on 13: 00 than that on 21: 00(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 1: 00 than that on any time point during the whole 24h in the LHb(P<0.05); The expression of Per2 was higher on 1: 00 than that on any time point during the whole 24h in the MHb(P<0.05); and c-fos mRNA and FOS protein in the MHb had no rhythmicity(P<0.05); The expression of c-fos mRNA and FOS protein was higher on 1: 00 than that on 13: 00 and 17: 00 in the MHb(P<0.05).
The expression of Per2 and c-fos mRNA and FOS protein in the LHb could be maintained the rhyhmicity with higher expression during the subjective night than day in DD cycle after SCN was lesioned: The expression was higher om 21: 00 and 1: 00 than that on 1: 00(P<0.05). The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned in the MHb(P>0.05).
Discussion:
The expression of Per2 and c-fos mRNA and FOS protein in the LHb and Hb showed the rhyhmicity with higher expression during the projective day than night, while there was no obvious rhythnicity in the MHb on different tine point in LD cycle. This rhythmic oscillation was coincident with the results from Prof. Zhao Hua who showed that cells in the LHb showed higher baseline firing rates during the day than during the night and cells in the MHb showed no rhythmic oscillation in vitro slice preparation. And in Zhao’s experiment of cells recorded in the LHb, 42% were tonically activated or suppressed by retinal illumination, while significantly fewer cells recorded in the MHb responded to retinal illumination(19%). It was reported that photosignal from the environment could project to the Hb directly and indirectly which made Hb receive the effect of photosignal. The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost after SCN was lesioned in the LHb and Hb and there was still no rhythmicity in the MHb. Therefore, we can conclude that SCN is very significant to the Hb to maintain its rhythmicity in LD cycle; And because the rhythmic expression of Per2 and c-fos mRNA and FOS protein oscillate in the LHb coincident to the Hb, the rhythmicity of the LHb master that of the whole Hb.
The expression of Per2 and c-fos mRNA and FOS protein showed the rhyhmicity with lower expression during the subjective day than the subjective night no matter in the LHb, MHb or in the Hb on different time point in DD cycle. Of all our results, there was no significance between 17: 00 and 21: 00(2 h before lights off and after lights off)about the expression of FOS protein which was identical with the story from Mahboubeh. Although Mahboubeh also found that the FOS protein expressed low on the subjective day and high on the subjective night in LHbM of hamster in DD cycle, the mechanism is excluded. The expression of Per2 and c-fos mRNA and FOS protein in the LHb could be maintained the rhyhmicity with higher expression during the subjective night than the subjective day in DD cycle after SCN was lesioned. Although the expressive difference abolished among some time points, the difference between 13: 00 and 1: 00 still existed in the LHb; The rhythmic expression of Per2 and c-fos mRNA and FOS protein lost in the MHb after SCN was lesioned. Thus, we can conclude that SCN is not indispensable to the Hb in DD; And the results futher support the phenomena: the rhythmicity of the LHb master that of the whole Hb.
Conclusion:
On the whole, there is a rhythmic oscillation in the LHb, MHb and Hb. Hb must rely on SCN in LD cycle, and SCN is very important. Although the expressive difference abolished among some time points in DD cycle, the expression of Per2 and c-fos mRNA and FOS protein still maintain the rhyhmicity with lower expression during day than the subjective night in the LHb, MHb or in the Hb. Hb can oscillates independent of SCN in DD cycle, but it is not a fully automic oscillation. Hb may participate the regulation of circadian rhythm system on the body and may play a different role from SCN in this system.
引文
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