Chinese Journal of Tissue Engineering Research ›› 2021, Vol. 25 ›› Issue (11): 1785-1790.doi: 10.3969/j.issn.2095-4344.3100
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Luo Yaxin, Bi Haoran, Chen Xiaoxu, Yang Kun
Received:
2020-07-17
Revised:
2020-07-21
Accepted:
2020-08-25
Online:
2021-04-18
Published:
2020-12-22
Contact:
Yang Kun, MD, Associate professor, Department of Periodontology, Stomatological Hospital Affiliated to Zunyi Medical University, Zunyi 563000, Guizhou Province, China
About author:
Luo Yaxin, Master candidate, Department of Periodontology, Stomatological Hospital Affiliated to Zunyi Medical University, Zunyi 563000, Guizhou Province, China
Supported by:
CLC Number:
Luo Yaxin, Bi Haoran, Chen Xiaoxu, Yang Kun. Extracellular matrix and tissue engineering regeneration and repair[J]. Chinese Journal of Tissue Engineering Research, 2021, 25(11): 1785-1790.
2.1 细胞外基质的定义 组织及器官都包含细胞和非细胞成分,彼此之间相互交错形成一个稳定且良好的复合结构。其中的非细胞成分即为细胞外基质,它是由细胞合成、分泌的生物大分子(如原纤维蛋白、蛋白聚糖、糖胺聚糖和糖蛋白等)在细胞表面或细胞之间构成的非细胞三维高分子网络[5],基质成分彼此之间及细胞与黏附受体结合形成一个复杂的网络,以便细胞驻留在该网络中的所有组织和器官中。细胞外基质分布于细胞间,尤其是结缔组织中细胞群体相对稀疏,其含量更为丰富[4],在为细胞提供生存所需的立体空间基础上,通过与细胞黏附分子、调节因子和连接蛋白等作用调控多种细胞功能,例如增殖、迁移、分化等[6-8],具有维持组织结构和参与生物反应的双重作用,对于维持正常的体内平衡至关重要。 2.2 细胞外基质的组成 细胞外基质可按其成分和结构的不同分为两种类型:间质和细胞周围基质[9-10],其中间质围绕细胞,而细胞周围基质则与细胞紧密接触,它们除了为细胞提供空间结构外,还可以进行相关信号转导、促进细胞内和细胞间的相互作用[11-12]。细胞外基质中富含胶原蛋白、层粘连蛋白、弹性蛋白、蛋白聚糖及各种细胞因子等[13],这些成分独立或协同与细胞相互作用发挥各自的功能。在受伤的皮肤中,层粘连蛋白可为皮肤细胞的黏附和生长提供底物,以覆盖伤口区域,重建完整的皮肤[14]。FERRUZZI等[15]发现弹性蛋白中的fibulin-5与心血管疾病存在一定相关性,缺乏fibulin-5的小鼠动脉压力明显增高,甚至会导致动脉功能丧失而诱发疾病。更有研究发现,基质细胞蛋白表达的改变与机体生理病理变化密切相关,它们在胚胎发育过程中表达最高,而在成人组织中表达逐渐减弱,但当机体发生例如炎症反应、伤口修复等活动时,基质细胞蛋白会大量激活以进行动态组织重塑和细胞运动[16]。此外,细胞外囊泡被认为是细胞外基质的重要组成部分,其包含多种表面分子,使它们能够靶向受体细胞并与细胞外基质相互作用,在细胞间转移化学信号来调节细胞行为[17-18]。 2.3 细胞外基质的提取方法 细胞外基质存在于细胞间,根据不同组织种类、细胞构成和体积密度可选用不同的脱细胞方法获取细胞外基质,目前常用的脱细胞方法可分为3大类:物理法、生物化学法及灌注脱细胞法[19]。物理法主要包括反复冻融、搅拌浸泡、高压力及超临界流体[20-21],其主要目的是通过直接机械力对细胞进行破坏。生物化学法包括pH值和酸碱度、非离子洗涤剂、离子洗涤剂、两性离子洗涤剂、酶制剂、螯合剂及毒素等[22-23]。此外,灌注脱细胞法也是提取方法之一[24],旨在保留器官的三维结构同时又能消除实质细胞群体。 2.4 细胞外基质的作用 2.4.1 命运指导 细胞外基质位于细胞周围,除了作为细胞生存及进行生理活动的场所,还扮演信号分子的角色,通过细胞膜表面受体传递信号对细胞增殖、迁移、分化等活动产生影响[25]。研究发现,细胞外基质可调节神经发育,与神经管和新皮质的形态发生有关[26-27],从而影响发育中的神经组织的形状。在对细胞外基质与增殖信号传导调节的研究中发现,细胞外基质中调节细胞增殖的关键成分(如蛋白聚糖、层粘连蛋白和整联蛋白)出现异常会导致细胞增殖能力减低,进而对组织的发育产生不利影响[28]。细胞迁移对于胚胎发生至关重要,而细胞迁移方式各有区别,既可单独活动又能以集体为单位进行迁移。HOCKMAN等[29]通过对神经嵴细胞的发育进行研究发现,细胞外基质可调节细胞迁移的方式,其主要原因可能与细胞外基质的内在成分改变有关。CHENG等[30]研究指出,细胞外基质成分与细胞迁移机制之间的相互作用非常复杂,通常需要各种膜结合和分泌性细胞蛋白在细胞外基质界面进行功能性相互作用,从而介导或调节胚胎细胞迁移。细胞外基质不仅可以介导细胞运动,同时还可维护组织器官的完整性,如在牙齿发育过程中对细胞的生物学指导和体内平衡起一定作用。LI等[31]对小型猪的乳牙牙胚细胞外基质成分进行研究,结果发现与牙齿发育相关的基因富集于细胞外基质中,这对于实现牙齿的完整发育十分重要。 2.4.2 信号传导 除了自身作为信号分子外,细胞外基质还调控信号分子(如生长因子和细胞因子)的转导,从而参与细胞与细胞之间的交流。细胞外基质是生长因子及细胞因子的重要来源及储存场所,二者会在特定条件下被释放,通过激活细胞信号而有助于细胞间信号传递并发挥功能[32]。组织器官的发育需要细胞外基质与细胞动态之间的协调平衡,细胞与细胞之间的黏附可介导组织和组织相互作用,细胞外基质结合的生长因子可以引起对组织发育和分化的深刻影响。转化生长因子β在调节胚胎发生、组织发育和干细胞命运方面起着重要作用,其与特定的细胞受体结合,并通过调节基因表达来激活细胞内信号传导,从而改变细胞和组织的行为[33]。在心脏相关研究中发现,细胞外基质通过调节细胞因子和生长因子反应进而激活成纤维细胞和血管细胞增殖分化,促进心脏组织的修复[34]。通过优化细胞外基质的机械特性可以增强人胚胎干细胞的中胚层分化。PRZYBYLA等[35]通过对细胞外基质中的生长因子进行调节,在高顺应性基质中培养人胚胎干细胞,其中胚层祖细胞的分化明显增强。细胞外基质是所有组织的生物活性成分,它不仅以细胞外基质-细胞方式参与细胞活动,更可以对细胞-细胞间信息交流进行调控,在指导细胞命运、影响组织发育和体内平衡方面发挥重要作用。 2.4.3 疾病诊断及治疗 随着人们对细胞外基质研究的深入,其作用被不断发掘,应用也越来越广泛。细胞外基质成分复杂多样并可触发多种生物学活动,对正常动态平衡至关重要,但是细胞外基质重塑也会导致许多疾病的发生,其中异常的分子是引发或发展疾病的关键因素[36-37],因此寻找它们会有助于疾病的诊断。如马凡氏综合征是一种由基因突变引起的常染色体显性遗传性结缔组织疾病,这种基因编码原纤维蛋白1,突变则会导致细胞外基质中原纤维蛋白1的数量减少或结构异常,从而极大地影响组织的结构和功能[38]。外泌体是膜结合的小囊泡,由正常细胞和癌细胞释放,它们被认为是细胞外基质不可或缺的组成部分。研究发现,外泌体整合素α6β4和α6β1与肿瘤肺转移有关,而外泌体整合素αvβ5与肿瘤肝转移有关,对它们进行调控后肺和肝的转移率出现下降趋势[39]。 除了对疾病的诊断有帮助,细胞外基质还可作为疾病药物治疗的靶点。研究表明当改变细胞外基质涂层配体的系链长度时,黏着力大小和细胞黏附强度会受到影响。用具有更长束缚长度的精氨酸-甘氨酸-天冬氨酸(Arg-Gly-Asp,RGD)配体涂覆坚硬的细胞外基质,会导致细胞感觉到更柔软的细胞外基质,从而控制癌症中机械转导介导的YAP/TAZ核定位,这可能是针对癌症的新颖有效治疗方法[40]。软骨和内皮下细胞外基质被越来越多地用于骨关节炎和类风湿关节炎的治疗中,被认为是治疗关节炎的靶标。胰岛素样生长因子1纳米复合物确保在软骨损伤的大鼠模型中保留了软骨完整性,并且纳米复合物治疗的动物的滑膜白细胞介素1β水平显著降低[41]。CHEN等[42]发现无活性相关肽通过基质金属蛋白酶2可裂解的接头与抗体的抗原结合Fv结构域融合,并且抗肿瘤坏死因子α或抗表皮生长因子抗体均引起配体结合减少,直到无活性相关肽介导的位阻被裂解和减弱,这些与无活性相关肽融合的前抗体在血清中稳定,表明有可能进行全身性给药并实现局部递送。 2.5 细胞外基质与再生修复 组织缺损再生修复一直是组织工程研究的核心,如何完善功能性修复更是成功的关键因素。理想的组织工程包括种子细胞、支架材料、器官构建和临床使用,其中支架材料和种子细胞是目前研究的重点内容。细胞外基质作为一种精密有序的网络结构能提供最接近于体内细胞生长的微环境,加之其富含的各种活性分子为细胞活动提供了基础,因此逐渐受到学者们的关注并应用于再生医学领域中。 2.5.1 软组织再生修复 各种急慢性原因均可导致软组织损伤,其中最常见的是皮肤软组织损伤。对于严重皮肤损伤的修复,确保足够的血管生成是首要因素。细胞外基质及其生物大分子(如胶原蛋白、弹性蛋白等)可通过诱导特定细胞行为、递送细胞或与特定因子及材料结合以促进血管再生。LAIVA等[43]研究发现胶原蛋白可与携带编码基质衍生因子1α结合来促进血管生成,是一种克服伤口愈合过程中不良血管反应的有效方法。MITHIEUX等[44]利用弹性蛋白构建一种名为HeaTro的支架材料,将成纤维细胞在HeaTro上培养一段时间后发现其增殖、迁移及黏附能力提高;在小鼠体内实验中,HeaTro植入组伤口愈合最佳,组织切片进一步证实了这一结果。脂肪组织是人体重要的组织结构之一,脂肪组织的功能随着年龄的增长会逐渐减退。当出现进行性脂肪营养不良、乳房切除和创伤性畸形引起的软组织损伤时,脂肪组织将延缓甚至丧失自我更新能力。KOKAI等[45]通过体外实验及动物模型体内植入实验认为细胞外基质可重塑和再生皮下脂肪软组织。神经系统主要由神经组织构成,分为中枢和周围神经系统两大部分。由于中枢神经系统的再生能力有限,成熟的神经元不能增殖分裂以替代损伤的细胞;并且受损伤组织内又产生了抑制神经细胞和轴突生长的环境,因此使其自身修复难以实现。SCHAEFFER等[46]发现细胞外基质在神经系统中具有特定调节功能,经深入研究后认为发挥这一作用的主要物质是细胞外基质的一种糖蛋白——Fibulin-2,它不仅可以参与神经胶质瘢痕部位细胞外基质的重塑,还可以影响星形胶质细胞的迁移活动,对中枢神经系统的修复和再生具有重要意义。牙龈是口腔软组织之一,牙龈退缩在牙周组织缺损中较为常见,侧向转位瓣术被认为是治疗牙龈退缩较为成功的方法。AHMEDBEYLI等[47]将脱细胞真皮基质移植物与侧向转位瓣术相结合对退缩牙龈进行修复,结果显示术后牙龈厚度及高度明显增高,缺损恢复较为理想,这将有利于牙周组织的健康和美观。 2.5.2 骨组织再生修复 组织再生中骨缺损的修复较为困难,自体骨移植虽是最佳选择,但由于来源匮乏、用量不足等原因而应用受限;异体骨因免疫排斥问题同样无法在临床上普及。细胞外基质来源广泛,种类丰富,并在干细胞分化中起重要作用,因而在骨组织工程中得以应用。ZHANG等[48]通过体外实验比较脂肪干细胞经人骨髓基质细胞来源细胞外基质培养前后的增殖及成骨分化能力,结果发现人骨髓基质细胞来源细胞外基质作用后脂肪干细胞的增殖速率较对照组提高,成骨集落及成骨标记表达得到改善,新生骨组织也更多。有学者将细胞外基质定植在钛基板上并用其培养骨髓间充质干细胞,发现该细胞外基质能够调节骨髓间充质干细胞的形态、增殖和成骨分化,认为细胞外基质在骨组织工程和其他相关生物医学中具备一定的应用潜力[49]。WEI等[50]通过体内外实验观察脂肪干细胞来源细胞外基质对骨髓间充质干细胞成骨分化的影响,体外实验结果显示,脂肪干细胞来源细胞外基质可促进骨髓间充质干细胞的成骨分化;将其混合聚(癸二酰基甘油二酯)支架植入大鼠颅骨缺损处,通过CT、荧光标记和组织学观察证实骨缺损修复效果显著增强。因此,脂肪干细胞来源细胞外基质可以促进骨髓间充质干细胞的成骨,并且脂肪干细胞来源细胞外基质和聚(癸二酰基甘油二酯)组合可以协同刺激骨形成,可能为骨骼再生提供更好的支持。 2.5.3 其他再生修复 由于心脏的极端结构和功能复杂性,心脏组织工程学面临着越来越高的需求,并且也面临着巨大的困难。心脏细胞外基质由于其复杂的天然结构,已被广泛应用于心脏组织工程。细胞外基质不仅通过其结构成分对心脏机械特性产生影响,而且还能通过调节细胞反应的分子信号来促进心脏的动态平衡,因此细胞外基质有可能在心脏组织再生修复中起关键作用[51]。肝脏是一种实质性器官,具有一定的自我再生能力,但当其发生严重病变(如晚期肝病、急性肝衰竭等)时受损的肝脏则无法自行恢复。细胞外基质可用作肝细胞培养的底物材料。AGARWAL等[52]对HepG2细胞构建含有细胞外基质的2D和3D培养环境,发现在2D培养中HepG2细胞的白蛋白、尿素、糖原和糖胺聚糖合成率提高,在含有细胞外基质的3D培养中HepG2细胞表现出高度分化和极化的表型,其成熟肝细胞标志物的表达增高,并且还形成胆小管样结构。这些结果表明细胞外基质在肝脏组织再生修复中具备优越性,可对开发用于组织工程的肝组织构建提供一定帮助。肾脏组织严重受损时通常伴有全身症状,极大影响患者的健康状况,甚至可能危及生命。相关文献报道,肾脏细胞外基质可为细胞的重建提供确切功能和结构环境,无论是用作细胞外基质支架还是生物打印都可能促进功能性肾脏的生成,在肾脏组织工程中至关重要[53]。先天性疾病、炎症和感染及手术损伤等多种因素可导致严重的小肠损伤,并伴有较高的发病率和死亡率。组织工程化的小肠已被广泛研究用于组织修复和重建,细胞外基质对小肠的修复作用也是毋庸置疑的[54]。 2.6 细胞外基质与组织工程 随着组织工程研究的兴起,结合细胞外基质自身的生物学优点,其在组织再生方面的应用也越来越广泛。通常,细胞外基质在组织再生中扮演细胞支架角色并表现出良好的再生修复能力。YANG 等[55]使用人骨髓间充质干细胞来源细胞外基质作为体外软骨细胞培养的底物,并检测其作为细胞支架用于软骨修复的适用性,发现人骨髓间充质干细胞来源细胞外基质无论在细胞培养还是在组织修复方面均展现出优势。脱细胞细胞外基质应用广泛。CHEN等[56]对斑马鱼心脏进行脱细胞处理得到脱细胞斑马鱼心脏来源细胞外基质,随后进行相关体内外实验,结果发现这种细胞外基质在体外可促进人心脏前体细胞的增殖和迁移,将其植入小鼠心肌梗死模型中能有效促进心肌组织再生,改善心脏功能。LIN等[57]制备猪坐骨神经脱细胞外基质,体外实验发现猪坐骨神经脱细胞外基质对许旺细胞增殖能力具有促进作用;在大鼠坐骨神经缺损模型中,猪坐骨神经脱细胞外基质组表现出的神经再生能力强于对照组,是一种有望用于周围神经再生的生物材料。此外,PATI等[58]采用多种方法处理软骨细胞外基质并结合水凝胶技术和3D打印技术制成支架材料,证明了脂肪干细胞可在该支架上分化形成软骨细胞并进一步形成软骨组织。 最近一些研究将细胞外基质更精细化处理,将它制成颗粒或者粉末单独或与其他材料联合应用于组织再生修复中,得到的结果似乎更理想。有学者通过一系列涉及粉碎、筛分和去细胞化过程得到细胞外基质颗粒,将其作用于骨髓干细胞发现细胞增殖率提高并在培养后期分化为成熟软骨细胞[59]。目前,粉末状细胞外基质已被用于将生长因子输送到难以直接进入的区域,例如肝脏。研究表明,肝源性细胞外基质粉末可携带生长因子直接输送到肝实质中,以诱导肝细胞增殖和肝再生[60]。此外,细胞外基质粉末具有良好的生物活性及生物相容性,在3D打印技术中联合使用有利于提高构建体的生物活性[61]。除了方法以外,材料的选择同样会对结果产生不同程度影响。有研究通过控制骨髓间充质干细胞在不同阶段的分化制备了干细胞、软骨形成、肥大和成骨阶段的4种细胞外基质支架,实验显示它们对骨髓间充质干细胞的成骨分化显示出不同的影响,其中肥大组细胞外基质支架对骨髓间充质干细胞成骨分化的促进作用最大,软骨形成组和成骨组居中,而干细胞组的促成骨分化能力最低[62]。因此,正确的方法协同最佳的材料才能使细胞外基质在组织工程中发挥最大作用。 "
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