Chinese Journal of Tissue Engineering Research ›› 2022, Vol. 26 ›› Issue (1): 125-129.doi: 10.12307/2022.021
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Wei Shan1, Li Qin2, Su Yi2
Received:
2020-12-14
Revised:
2021-01-05
Accepted:
2021-02-05
Online:
2022-01-08
Published:
2021-10-25
Contact:
Su Yi, Master, Associate chief physician, Department of Stomatology, Eye & ENT Hospital of Fudan University, Shanghai 200031, China
About author:
Wei Shan, Lecturer, Huizhou Health Vocational and Technical College, Huizhou 516025, Guangdong Province, China
Supported by:
CLC Number:
Wei Shan, Li Qin, Su Yi. Research hotspots and application potential of stem cells from human exfoliated deciduous teeth[J]. Chinese Journal of Tissue Engineering Research, 2022, 26(1): 125-129.
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2.1 乳牙牙髓干细胞的一般生物学特性 2.1.1 乳牙牙髓干细胞的来源、一般特征 乳牙牙髓干细胞的胚胎发育起源于神经嵴外胚层,这些组织迁徙到颌面部后形成外胚间充质,MIURA等[1]发现体外培养的乳牙牙髓干细胞多位于牙髓血管周围,提示乳牙牙髓干细胞可能起源于管周微环境。乳牙牙髓干细胞大多呈长梭形,成纤维细胞样,少数为多角形和椭圆形,细胞体积小、核大、着色深,卵圆形,有一个或几个核体,胞浆着色浅。 2.1.2 乳牙牙髓干细胞的衰老与凋亡 细胞衰老的标志包括细胞生长停滞、DNA损伤灶和衰老相关半乳糖苷酶的表达。研究表明,在牙髓干细胞增殖的同时,也同样发生着细胞衰老,并且衰老细胞的分化潜能明显降低,还会分泌对邻近组织有害的因子。现已知与营养感知路径相关的5'腺苷单磷酸活化蛋白激酶(AMPK)途径是调节细胞衰老的因素之一[2]。 细胞凋亡一般有3种典型通路,包括外源性通路、内源性通路和颗粒酶通路。Caspase属于半胱氨酸蛋白酶,这些蛋白酶是引起细胞凋亡的关键酶,一旦被信号途径激活,能将细胞内的蛋白质降解,使细胞不可逆的走向死亡。例如Caspase-8、Caspase-9和Caspase-10接收到信号后,能通过自剪接而激活,然后引起caspase级联反应,依次激活Caspase-3、Caspase-6、Caspase-7。QIAN等[3]通过定量PCR以及蛋白印迹法证实了牙髓干细胞中细胞色素C的表达以及线粒体介导的凋亡通路活动,并发现与成人牙髓干细胞相比,乳牙牙髓干细胞的细胞凋亡显著增加。虽然Caspase-8和Caspase-9的表达在乳牙内高于成人恒牙,但只有通过RNA干扰敲除牙髓干细胞中的Caspase-9,细胞凋亡、Caspase-3的表达和活性才会明显减少。因此,研究认为Caspase-9调节乳牙牙髓干细胞的凋亡。YAMAZA等[4]研究发现高浓度的游离胆红素通过线粒体途径诱导乳牙牙髓干细胞的凋亡,这可能与丝氨酸/苏氨酸激酶(AKT)信号通路和细胞外信号相关激酶1和2 (ERK1/2)信号通路的抑制以及核转录因子κB (NF-κB)信号通路的激活相关。 2.1.3 乳牙牙髓干细胞的微环境 干细胞的微环境是指通过细胞之间的特定相互作用和多种生物活性分子来调节干细 胞/祖细胞的增殖、生存、迁移和衰老。乳牙牙髓干细胞的微环境在出生前就形成了,并一直维持到恒牙萌出前。在这一短暂时期,机体生长活跃,并维持微环境中的干细胞活力,而这些微环境并没有严重受到遗传和/或环境因素的影响[5]。因此,与从恒牙分离的牙髓干细胞相比,从乳牙中分离的牙髓干细胞为未来的细胞疗法提供了理想的细胞来源。 碱性成纤维细胞生长因子维持乳牙牙髓干细胞的干性,并调控乳牙牙髓干细胞的成牙分化以及矿化。NOWWAROTE等[6]在体外乳牙牙髓干细胞的成骨/成牙分化过程中添加10 μg/L碱性成纤维细胞生长因子,发现碱性磷酸酶mRNA表达和碱性磷酸酶活性降低,ANKH mRNA表达增加,矿物质沉积减少。加入外源性无机焦磷酸盐可抑制矿化,增加ANKH、COL1A1和SPP1 mRNA的表达,而加入外源性无机磷酸盐可增加矿化和成骨细胞特异性转录因子OSX、ANKH、SPP1、DMP1 mRNA的表达。NOWWAROTE等[6]研究首次揭示了碱性成纤维细胞生长因子对乳牙牙髓干细胞中无机磷酸盐/无机焦磷酸盐调节因子的影响。汪璐璐等[7]研究认为在乳牙生理性根吸收过程中α7亚型烟碱型乙酰胆碱受体(α7nAChR)表达上调,参与对核因子κB受体活化因子/骨保护素(RANKL/OPG)的调节,影响乳牙牙髓干细胞的破骨分化能力。张灵芝[8]研究发现肿瘤坏死因子刺激下人乳牙牙髓干细胞的破骨相关基因CTSK和TRAP mRNA表达量明显上升,差异有显著性意义,因此认为炎症微环境可以导致人乳牙牙髓干细胞促破骨细胞形成能力增强,可能导致乳牙早失。 2.1.4 端粒长度 在大多数成体细胞中,细胞分裂和DNA复制是端粒缩短的结果。增殖活跃的干细胞端粒长度为10- 20 kb,而在体细胞中细胞端粒长度为5-15 kb。 2.2 乳牙牙髓干细胞的各项功能 2.2.1 乳牙牙髓干细胞的增殖功能 研究显示,乳牙牙髓干细胞具有较高的增殖率,每颗切牙有12-20个细胞能形成克隆集落并高度增殖。NAKAMURA等[9]将乳牙牙髓干细胞、成人牙髓干细胞和骨髓间充质干细胞进行比较,发现乳牙牙髓干细胞比成人牙髓干细胞和骨髓间充质干细胞的增殖能力更高。Real-time PCR显示,乳牙牙髓干细胞中成纤维细胞生长因子2、转化生长因子β2、Ⅰ型胶原和Ⅱ型胶原的表达显著高于骨髓间充质干细胞和成人牙髓干细胞。KUNIMATSU 等[10]研究得出类似的结论,与骨髓间充质干细胞和成人牙髓干细胞相比,乳牙牙髓干细胞拥有更高的增殖能力,以及更高的碱性成纤维细胞生长因子和骨形态发生蛋白的基因表达。 ALMEIDE-JUNIOR等[11]研究认为低剂量的激光照射对乳牙牙髓干细胞的增殖也有影响,实验发现用2次或3次 2.5 J/cm2红色激光照射乳牙牙髓干细胞,在48 h内拥有更高的细胞存活率;而单次2.5 J/cm2照射时,在48 h内不会刺激细胞代谢活动,在72 h内也不会刺激细胞增殖,单次照射剂量为5.0 J/cm2和 7.5 J/cm2以及应用3次2.5 J/cm2可以在72 h维持细胞活性,并刺激乳牙牙髓干细胞增殖。丁钰等[12]研究发现富血小板血浆对乳牙牙髓干细胞的增殖和向成骨分化均具有一定的促进作用,随着浓度的增大和作用时间的延长,其促进增殖的能力也随之增强。 2.2.2 乳牙牙髓干细胞的多向分化功能 (1)成牙本质/牙髓分化:SAKAI等[13]发现体外诱导乳牙牙髓干细胞表达血管内皮因子受体2、CD31和血管内皮钙粘蛋白,分化为血管内皮细胞;把乳牙牙髓干细胞和生物支架接种在牙片上并将其植入裸鼠皮下,发现乳牙牙髓干细胞可以分化为能产生管状牙本质的成牙本质细胞和血管内皮细胞。CORDEIRO等[14]将乳牙牙髓干细胞和皮肤血管内皮细胞接种在可降解的生物支架上,并将其植入裸鼠皮下,观察到乳牙牙髓干细胞最终形成牙髓样组织,证实在一定条件下,乳牙牙髓干细胞可以被诱导成为牙髓组织。 CASAGRANDE等[15]发现牙本质来源骨形态发生蛋白2可以诱导乳牙牙髓干细胞向牙本质细胞分化,表达成牙本质细胞的表面标志,而阻断骨形态发生蛋白2信号通路乳牙牙髓干细胞向成牙本质细胞分化受到抑制,得出结论:乳牙牙髓干细胞向成牙本质细胞分化需要骨形态发生蛋白2参与。 (2)向成骨方向分化:研究证明,牙髓干细胞在特定的细胞因子诱导下,可以向成骨细胞分化。骨涎蛋白、骨桥蛋白都被研究证明可以诱导乳牙牙髓干细胞选择性分化为成骨细胞,并且可促进成骨细胞的黏附和分化,同时是早期成骨分化的标志物。CHADIPRIALLA等[16]发现,视黄酸和胰岛素共同诱导乳牙牙髓干细胞向成骨方向分化效果最好。SU等[17]研究表明,锶可以成为乳牙牙髓干细胞成骨的有效诱导剂,乳牙牙髓干细胞被包封在含有磷酸锶的水凝胶中,成骨相关基因和骨粘连蛋白的表达水平均上调。 MIURA等[1]认为乳牙牙髓干细胞可以引导新骨形成,但不能直接分化成骨细胞,WANG等[18]则认为乳牙牙髓干细胞具有成骨潜能,在成骨诱导后2周乳牙牙髓干细胞能分化成功能性的类成骨细胞。 (3)向神经方向分化:MIURA等[1]将乳牙牙髓干细胞在成神经条件下培养后,持续表达神经胶质细胞的标志物。MORSCZECK等[19]比较乳牙牙髓干细胞、人牙髓干细胞向神经方向分化情况,在特定条件下,乳牙牙髓干细胞向神经分化的时间比人牙髓干细胞快4倍左右,推测乳牙牙髓干细胞有更好的神经细胞分化潜能。YANG等[20]将乳牙牙髓干细胞分别转染Noggin、Y-27632以及Noggin和Y-27632的结合体,发现转染Y-27632后,乳牙牙髓干细胞活力增强,形成细胞集落和神经球,说明Y-27632可以促进乳牙牙髓干细胞的增殖。实验还观察到,Noggin结合Y-27632处理的乳牙牙髓干细胞呈现出典型的神经元样细胞形态和网状突起。无论是单独Noggin或Y-27632转染还是Noggin、Y-27632两者一起转染,乳牙牙髓干细胞中NSE、Nestin和GFAP水平均明显升高,Noggin、Y-27632两者一起转染的乳牙牙髓干细胞中NSE、Nestin和GFAP水平最高,说明二者对乳牙牙髓干细胞向神经元样细胞分化具有协同作用。 (4)向成脂方向分化:2003年,MIURA等[1]发现乳牙牙髓干细胞在成脂条件下诱导培养5周后,半定量RT-PCR检测到脂肪细胞特有的转录因子表达上调,说明乳牙牙髓干细胞存在成脂方向分化的可能。ZHANG等[21]将乳牙牙髓干细胞用成脂分化培养基诱导培养后产生大量脂质空泡,脂蛋白酶增加到对照细胞的31.7倍,证实乳牙牙髓干细胞在诱导条件下能分化成脂肪细胞。还有一些研究也用油红O染色证实了乳牙牙髓干细胞具有成脂分化的潜能[22]。不过,PIVORIUūNAS 等[23]研究却认为乳牙牙髓干细胞并没有太多的成脂分化能力。 (5)向其他方向分化:ANNIBALI等[24]研究发现乳牙牙髓干细胞在体外可以诱导分化为软骨样细胞。ISHKITIEV等[25]将分离培养的乳牙牙髓干细胞在含肝细胞生长因子的培养基中进行诱导培养,发现其可表达多种肝细胞特异性标志物。KIM等[26]证实了乳牙牙髓干细胞诱导后具有血管周围细胞的特征并通过定量PCR发现乳牙牙髓干细胞能够表达血管周细胞标志物。PAINO等[27]将牙髓干细胞培养于改良DMEM培养基中,发现培养出的细胞表达高水平的血管内皮生长因子等,证实乳牙牙髓干细胞具有向血管方向分化的潜能。GOVINDASAMY等[28]研究报道了从完整乳磨牙中分离出的乳牙牙髓干细胞具有分化成胰腺样细胞的能力。 2.3 乳牙牙髓干细胞的临床治疗研究 2.3.1 乳牙牙髓干细胞在口腔疾病方面的研究 (1)在牙周骨缺损中的应用:WANG等[29]研究发现乳牙牙髓干细胞外泌体对牙周膜干细胞活力没有细胞毒性;茜素红染色、碱性磷酸酶活性以及成骨基因(RUNX2、OPN、OCN)表达上调都表明乳牙牙髓干细胞外泌体促进了牙周膜干细胞的成骨分化。研究进一步发现提高Smad1/5/8的磷酸化作用和增加细胞核内β-连环蛋白的表达可以激活BMP/Smad和Wnt/β-连环蛋白信号通路。用特定的抑制剂(豆蔻素和LDN193189)抑制这两个信号通路,其成骨分化能力明显减弱。在乳牙牙髓干细胞和乳牙牙髓干细胞外泌体中Wnt3a和BMP2表达明显上调,因此这个研究说明乳牙牙髓干细胞外泌体通过Wnt3a和BMP2促进牙周膜干细胞的成骨分化,这也为乳牙牙髓干细胞外泌体治疗牙周炎导致的骨缺损提供了新见解。 (2)在口腔黏膜疾病中的应用:DU等[30]将乳牙牙髓干细胞注入舍格仑综合征小鼠模型体内,研究发现唾液分泌增加,下颌下腺的炎症反应减轻,血清、下颌下腺,唾液中的炎性细胞因子减少,凋亡细胞减少,下颌下腺中ATG5和苄氯素1的表达减少,并对其他器官无影响;注射的干细胞迁移至脾脏和肝脏,而不是下颌下腺。研究认为乳牙牙髓干细胞可以通过减少炎症细胞因子,调节炎症微环境,减少细胞凋亡和自噬来减轻舍格仑综合征小鼠唾液过少的症状。 (3)在颌面部骨缺损中的应用:ZHENG等[31]将小型猪自体乳牙牙髓干细胞植入β-磷酸三钙支架培养7 d,然后再将其移植到猪下颌骨缺损部位,6个月后缺损部位新生骨成功修复下颌骨缺损。YAMADA等[32]从幼犬乳牙中提取干细胞,植入母犬下颌骨内,研究发现乳牙牙髓干细胞具有成骨能力,使子女和父母之间的移植成为可能。HIRAKIA等[33]研究发现乳牙牙髓干细胞对唇腭裂和骨重建可发挥较好效果。PEREIRA等[34]发现乳牙牙髓干细胞可以促进面神经损伤后的重建。 (4)在牙髓再生中的应用:ROSA等[35]在体外将乳牙牙髓干细胞混合重组人Ⅰ型胶原并注入人前磨牙,可以观察到牙髓样组织的出现,并产生新的贯穿根管的管状牙本质,提示乳牙牙髓干细胞在牙髓组织工程中的潜在应用。 2.3.2 乳牙牙髓干细胞在其他疾病方面的研究 (1)脊髓损伤:2012年,SAKAI等[36]发现乳牙牙髓干细胞比骨髓间充质细胞拥有更高的神经再生活性,将其移植入脊髓损伤大鼠模型后,改善了大鼠后肢运动功能;乳牙牙髓干细胞可以抑制脊髓损伤大鼠少突胶质细胞和神经元的凋亡。NICOLA等[37]将乳牙牙髓干细胞移植联合跑步训练治疗实验性脊髓损伤大鼠,结果表明,乳牙牙髓干细胞存在神经保护以及抗炎作用;特别是,乳牙牙髓干细胞可以促进功能恢复以及增加创伤后病灶附近的神经丝密度。这些结果充分证明乳牙牙髓干细胞在脊髓损伤后可以多方面利于其功能恢复。 (2)红斑狼疮:红斑狼疮是一种累及多脏器的自身免疫性疾病,YAMAZA等[38]将乳牙牙髓干细胞移植到系统性红斑狼疮小鼠体内,发现与骨髓间充质干细胞相比,乳牙牙髓干细胞的免疫调节作用更强。 (3)外伤性脑损伤:LI等[39]第一次证明了乳牙牙髓干细胞来源外泌体对大鼠外伤性脑损伤有治疗效果,至少部分是通过改变小胶质细胞的极化来减少神经炎症。 (4)色素性视网膜炎:LI等[40]发现乳牙牙髓干细胞和乳牙牙髓干细胞条件培养基对大鼠色素性视网膜炎都有治疗作用,增强视网膜神经纤维层的电活动,对视网膜光感受器层损伤的修复有明显促进作用,主要与乳牙牙髓干细胞的抗凋亡活性有关。因此认为乳牙牙髓干细胞是治疗视网膜变性的非常有前途的干细胞来源。 (5)肝疾病:ISHKITIEV等[25]将乳牙牙髓干细胞分化的肝细胞注射到肝硬化大鼠肝脏中,明显改善了大鼠的肝功能并提高了存活率。TAKAHASHI等[41]发现乳牙牙髓干细胞转化肝样细胞微球改善了肝功能,有抗纤维化作用。 (6)糖尿病:RAO等[42]认为乳牙牙髓干细胞为2型糖尿病提供了一种潜在的治疗手段。将乳牙牙髓干细胞注射入糖尿病大鼠体内,注射8周后,大鼠血糖水平显著降低,另外组织学分析胰岛和肝损伤明显减轻。 (7)其他疾病:由于乳牙牙髓干细胞的多分化性,使其在全身的各项疾病中都可能发挥作用,例如有学者发现乳牙牙髓干细胞能治疗急性肾损伤,它的治疗作用主要来自于其抗炎作用、抗细胞衰亡作用和旁分泌作用[43]。还有研究发现乳牙牙髓干细胞对肌萎缩、自身免疫性脑脊髓炎有治疗 效果[5,44]。 "
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