基于生物矿化提高生物耐热性的新策略
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摘要
生物矿化是指生物体内矿物形成的过程,它将自然界中的有机软物质和无机硬物质联系起来。在自然界中,大多数生命体都进化出了巧妙的生物矿化调控机制、能够构建出各种功能生物材料。例如,小至趋磁细菌中的纳米磁性微晶,还有介观的硅藻壳、植物硅晶石,以及大到宏观尺度的骨骼和牙齿等。特别需要提到的是,从海绵到温泉细菌,从低等硅藻到高等植物如水稻等都可以通过生物矿物来增强它们对抗恶劣自然环境的能力。
受生物矿化现象的启发,我们提出通过仿生矿化来改善生物体系在恶劣环境中的抗逆性。近年来,研究发现有机分子特别是蛋白质在生物矿化中起着至关重要的调控作用,科学家们能够利用功能性高分子或仿生多肽来模拟矿化蛋白控制无机矿化的过程,从而能够在近生理的温和条件下将无机功能矿物引入生物体中并赋予它们新的特性。酵母作为一种单细胞真核生物是实现该设想的良好模型。病毒疫苗在疾病预防和治疗中具有重要的地位,因此对它们进行生物功能化具有十分重大的科学和社会意义。本论文通过生物矿化手段分别在酵母和病毒疫苗表面引入无机矿物,并进一步关注材料功能化后的酵母或疫苗在热稳定性方面的提升。全文共分为六章：
第一章绪论是对生物矿化的背景及本论文研究线路进行概述。首先介绍了生物矿物的种类并对二氧化硅、磷酸钙、碳酸钙以及无定形矿物相进行了重点叙述；然后综述了蛋白、高分子和仿生多肽等有机分子在生物矿化中的调控作用,突出了生物体系对磷酸钙和二氧化硅矿化的调控；接着归纳了仿生矿化研究中的一些基本原理及相关应用；随后提出基于仿生矿化的生物功能化设想并进而明确了在疫苗矿化改造方面展开研究,希望获得热稳定良好的液态疫苗,同时也介绍现有提升疫苗热稳定的一些方法；最后提出本研究的思路和目标等。
第二章我们利用酵母细胞作为模型体系探索仿生矿化这种策略在生物功能化方面的应用。我们通过层层自组装并结合生物矿化手段对细胞进行了二氧化硅壳化修饰,研究了矿物壳对生物体的影响,特别关注细胞耐热性的提升。实验结果显示了人工引入的硅矿物层可以改进酵母细胞的耐热性,其原理是二氧化硅矿物层可以通过大量的氢键来禁锢细胞周围的水分子进而降低了热运动对细胞膜和细胞器的破坏。
第三章起我们将注意力集中在热稳定性疫苗的开发。该章以乙型脑炎病毒疫苗株(JEV SA14-14-2)为模型,利用原位矿化成功地在疫苗表面引入了一层纳米磷酸钙外壳。该外壳不影响疫苗基本生物学功能,但能够显著提高疫苗的热稳定性。该研究第一次提出了生物矿化策略可以用于制造出新型液态热稳定疫苗,为常温疫苗的研发提供了全新的思路。这种便宜、方便的制备热稳定疫苗的策略有利于免疫接种项目在发展中国家和贫困地区的推广
第四章中我们受生物体利用蛋白实现自矿化这一现象启发,通过反向遗传学手段获得了一株具有自矿化能力的基因工程疫苗。该疫苗可以在近生理环境下自发地矿化形成纳米磷酸钙外壳,矿化后的疫苗在保持良好的免疫原性的基础上也表现出显著增强的热稳定性。由于这种自矿化能力具有可遗传性,所以该方法突破了疫苗本身矿化能力的限制,为热稳定疫苗提供了一种普适性的制备方法,特别有利于热稳定疫苗的规模化生产。
第五章中我们的目标是通过改变矿化材料来提升矿化策略在制备热稳定疫苗方面的应用。我们注意到,自然界中温泉细菌和热带植物往往通过富集无定形二氧化硅来增强其耐热性。受此启迪,我们通过原位硅矿化在病毒疫苗表面引入无定形硅纳米簇,大幅度提升疫苗的热稳定性。例如,硅矿化后的肠道病毒71型和脊髓灰质炎病毒疫苗的热稳定性提高了7-10倍(磷酸钙矿化提高疫苗热稳定性仅3倍左右)。硅矿化改造疫苗在室温环境的存放可长达一个多月。进一步研究发现,这些无定形硅纳米簇通过表面羟基与周围的水分子形成大量的氢键,进而在疫苗表面形成水合硅层,降低了疫苗与周围水环境的化学键交换,提高了疫苗的热稳定性。基于该理解,我们提出了热稳定的硅锚新概念,也在化学原理上合理的解释了自然界中的生物体往往选择无定形硅材料作为抗热材料。
第六章我们对本论文的研究做了简要的归纳和总结,明确展示了生物矿化策略在改善生物耐热性、尤其是在制备热稳定疫苗方面的巨大前景。我们还进一步讨论了今后如何更好地发展和应用这种仿生策略实现基于材料科学的生物功能化：
Biomineralization refers to the processes by which living organisms deposit minerals, and it links the soft organic material and the hard inorganic materials on the earth. Along with the evolution, almost all organisms have evolved complicate and delicate biomineralization mechanisms to control the biomineral deposition, For example the magnetotactic bacteria deposit iron oxide in enclosed organic stealths, diatoms and sponges control the formation of the macroscopic silica structures, mollusks form calcite crystals in their shells, vertebrates generate apaptite in bones and enamels. Generally, organic macromolecules such as protein and glycoprotein play an important role in regulating these biomineralization processes. The polymers with similar residues and biomimetic polypeptides are the favorable substitution in the biomimetic studies, as they are easier to be obtained and controlled. Recently, they have been broadly adopted in the material synthesis and organism biomineralization.
Inspired by these biomineralization phenomena, functional inorganic materials can be introduced to living organisms in a biomimetic way to improve their performance in harsh environments. Organic biomolecules, such as protein and glycoprotein, play a key role in the control and regulation of biomineralization. Due to the complex of proteins, biomimetic polypeptides are widely used in preparing functional materials and studying organism mineralization. By introducing an inorganic exterior on organisms, we can confer them some new characteristics, such as themostability and water retention capability. Yeast, as a typical unicellular eukaryote, is a good living model in the research. Virus, which widely be used in material construction, cargo delivery and vaccine development due to its composition and structure simplicity, is also a good model. Futhermore, virus can feasibly be operated to display nucleating peptides by genetic engineering because viral genome is simple.
In this dissertation, we confer the organisms such as yeast and virus vaccine an inorganic exterior by biomimetic mineralization under mild condition, which significantly improved the organisms'thermostabiltiy and water retention capability. This study mainly focuses on the development of liquid thermostable vaccines by using a biomineralization strategy. This dissertation can be divided into five chapters: Chapter1. The introduction mainly composed of an overview of biomineralization, biogenic minerals such as silica, calcium phosphate (CaP), calcium carbonate, iron minerals and amorphous phrase; the roles of protein, polymer and peptide in the control of calcium phosphate and silica biomineralization; biomimetic principle and applications; yeast cell encapsulation and mineralization, cell surface modification with polymers and inorganic materials; the chemical and biological modification of viral capsid, and functional nanomaterials by using virus as the template; the importance of vaccines and their thermostability, approaches to thermostable vaccines; the strategy and aims of this study.
Chapter2. We conferred the yeast cell an inorganic calcium phosphate or silica shell by layer-by-layer method, and further studied the implications of this modification. The silica nanocoat could improve the thermostability and water retention capability upon drying condition.
Chapter3. A eggshell-like calcium phopahte shell was introduced on Japanese encephalitis virus by in situ biomineralization. This eggshell-like exterior significantly improved the thermostability of virus vaccine without impairing its biological characteristics. This study provides a novel approach to thermostable vaccine development in developing countries.
Chapter4. Using the human enterovirus type71(EV71) vaccine strain as a model, we suggest a combined, rational design approach to improve the thermostability and immunogenicity of live vaccines by self-biomineralization. The biomimetic peptides are rationally integrated onto the capsid of EV71by reverse genetics so that calcium phosphate mineralization can be biologically induced onto vaccine surfaces under physiological conditions, generating a mineral exterior. This engineered self-biomineralized virus was characterized in detail for its unique structural, virological and chemical properties. Analogous to many exteriors, the mineral coating confers some new properties on enclosed vaccines. Such a combination of genetic technology and biomineralization provides an economic solution for current vaccination programs, especially in developing countries that lack expensive refrigeration infrastructures.
Chapter5. We showed a feasible strategy to produce thermostable liquid vaccines by introducing a protective hydrated amorphous silica exteriors to them with tunable in situ silicification. The hydrated amorphous silica exterior does not impair the original biological behaviors of virus vaccines, but confers a significantly improved thermostability on them at a wide range of temperatures. After silicification, the storage period of Poliovirus and EV71virus at room temperature can be extended by about7-fold and10-fold respectively, realizing the storage of these vaccines at room temperature for more than one month. Silica exteriors confined nearby water molecules through hydrogen bonds formed between water and hydroxyl groups, and therefore decreased the exchange of ionic and hydrogen bonding between solution and interior vaccine. The technology may pose great impact on current vaccination program by providing an economic strategy for producing liquid thermostable vaccines. Based on these results, we suggest a new concept that the silica achors can improve organisms'thermostability. The study also provides a reasonable explanation why the nature organisms often choose amorphous silica as the biominerals.
Chapter6. We briefly summaried the studies in this dissertation. Our studies clearly demostrate that the biomimetic mineralization strategy can be successfully applied in improving organisms' themostability, and hold great promise in the development of liquid thermostable vaccines. We further disscussed how to better applied this functional materials based biomimetic strategy to improve the biologicals' applications in future.
受生物矿化现象的启发,我们提出通过仿生矿化来改善生物体系在恶劣环境中的抗逆性。近年来,研究发现有机分子特别是蛋白质在生物矿化中起着至关重要的调控作用,科学家们能够利用功能性高分子或仿生多肽来模拟矿化蛋白控制无机矿化的过程,从而能够在近生理的温和条件下将无机功能矿物引入生物体中并赋予它们新的特性。酵母作为一种单细胞真核生物是实现该设想的良好模型。病毒疫苗在疾病预防和治疗中具有重要的地位,因此对它们进行生物功能化具有十分重大的科学和社会意义。本论文通过生物矿化手段分别在酵母和病毒疫苗表面引入无机矿物,并进一步关注材料功能化后的酵母或疫苗在热稳定性方面的提升。全文共分为六章：
第一章绪论是对生物矿化的背景及本论文研究线路进行概述。首先介绍了生物矿物的种类并对二氧化硅、磷酸钙、碳酸钙以及无定形矿物相进行了重点叙述；然后综述了蛋白、高分子和仿生多肽等有机分子在生物矿化中的调控作用,突出了生物体系对磷酸钙和二氧化硅矿化的调控；接着归纳了仿生矿化研究中的一些基本原理及相关应用；随后提出基于仿生矿化的生物功能化设想并进而明确了在疫苗矿化改造方面展开研究,希望获得热稳定良好的液态疫苗,同时也介绍现有提升疫苗热稳定的一些方法；最后提出本研究的思路和目标等。
第二章我们利用酵母细胞作为模型体系探索仿生矿化这种策略在生物功能化方面的应用。我们通过层层自组装并结合生物矿化手段对细胞进行了二氧化硅壳化修饰,研究了矿物壳对生物体的影响,特别关注细胞耐热性的提升。实验结果显示了人工引入的硅矿物层可以改进酵母细胞的耐热性,其原理是二氧化硅矿物层可以通过大量的氢键来禁锢细胞周围的水分子进而降低了热运动对细胞膜和细胞器的破坏。
第三章起我们将注意力集中在热稳定性疫苗的开发。该章以乙型脑炎病毒疫苗株(JEV SA14-14-2)为模型,利用原位矿化成功地在疫苗表面引入了一层纳米磷酸钙外壳。该外壳不影响疫苗基本生物学功能,但能够显著提高疫苗的热稳定性。该研究第一次提出了生物矿化策略可以用于制造出新型液态热稳定疫苗,为常温疫苗的研发提供了全新的思路。这种便宜、方便的制备热稳定疫苗的策略有利于免疫接种项目在发展中国家和贫困地区的推广
第四章中我们受生物体利用蛋白实现自矿化这一现象启发,通过反向遗传学手段获得了一株具有自矿化能力的基因工程疫苗。该疫苗可以在近生理环境下自发地矿化形成纳米磷酸钙外壳,矿化后的疫苗在保持良好的免疫原性的基础上也表现出显著增强的热稳定性。由于这种自矿化能力具有可遗传性,所以该方法突破了疫苗本身矿化能力的限制,为热稳定疫苗提供了一种普适性的制备方法,特别有利于热稳定疫苗的规模化生产。
第五章中我们的目标是通过改变矿化材料来提升矿化策略在制备热稳定疫苗方面的应用。我们注意到,自然界中温泉细菌和热带植物往往通过富集无定形二氧化硅来增强其耐热性。受此启迪,我们通过原位硅矿化在病毒疫苗表面引入无定形硅纳米簇,大幅度提升疫苗的热稳定性。例如,硅矿化后的肠道病毒71型和脊髓灰质炎病毒疫苗的热稳定性提高了7-10倍(磷酸钙矿化提高疫苗热稳定性仅3倍左右)。硅矿化改造疫苗在室温环境的存放可长达一个多月。进一步研究发现,这些无定形硅纳米簇通过表面羟基与周围的水分子形成大量的氢键,进而在疫苗表面形成水合硅层,降低了疫苗与周围水环境的化学键交换,提高了疫苗的热稳定性。基于该理解,我们提出了热稳定的硅锚新概念,也在化学原理上合理的解释了自然界中的生物体往往选择无定形硅材料作为抗热材料。
第六章我们对本论文的研究做了简要的归纳和总结,明确展示了生物矿化策略在改善生物耐热性、尤其是在制备热稳定疫苗方面的巨大前景。我们还进一步讨论了今后如何更好地发展和应用这种仿生策略实现基于材料科学的生物功能化：
Biomineralization refers to the processes by which living organisms deposit minerals, and it links the soft organic material and the hard inorganic materials on the earth. Along with the evolution, almost all organisms have evolved complicate and delicate biomineralization mechanisms to control the biomineral deposition, For example the magnetotactic bacteria deposit iron oxide in enclosed organic stealths, diatoms and sponges control the formation of the macroscopic silica structures, mollusks form calcite crystals in their shells, vertebrates generate apaptite in bones and enamels. Generally, organic macromolecules such as protein and glycoprotein play an important role in regulating these biomineralization processes. The polymers with similar residues and biomimetic polypeptides are the favorable substitution in the biomimetic studies, as they are easier to be obtained and controlled. Recently, they have been broadly adopted in the material synthesis and organism biomineralization.
Inspired by these biomineralization phenomena, functional inorganic materials can be introduced to living organisms in a biomimetic way to improve their performance in harsh environments. Organic biomolecules, such as protein and glycoprotein, play a key role in the control and regulation of biomineralization. Due to the complex of proteins, biomimetic polypeptides are widely used in preparing functional materials and studying organism mineralization. By introducing an inorganic exterior on organisms, we can confer them some new characteristics, such as themostability and water retention capability. Yeast, as a typical unicellular eukaryote, is a good living model in the research. Virus, which widely be used in material construction, cargo delivery and vaccine development due to its composition and structure simplicity, is also a good model. Futhermore, virus can feasibly be operated to display nucleating peptides by genetic engineering because viral genome is simple.
In this dissertation, we confer the organisms such as yeast and virus vaccine an inorganic exterior by biomimetic mineralization under mild condition, which significantly improved the organisms'thermostabiltiy and water retention capability. This study mainly focuses on the development of liquid thermostable vaccines by using a biomineralization strategy. This dissertation can be divided into five chapters: Chapter1. The introduction mainly composed of an overview of biomineralization, biogenic minerals such as silica, calcium phosphate (CaP), calcium carbonate, iron minerals and amorphous phrase; the roles of protein, polymer and peptide in the control of calcium phosphate and silica biomineralization; biomimetic principle and applications; yeast cell encapsulation and mineralization, cell surface modification with polymers and inorganic materials; the chemical and biological modification of viral capsid, and functional nanomaterials by using virus as the template; the importance of vaccines and their thermostability, approaches to thermostable vaccines; the strategy and aims of this study.
Chapter2. We conferred the yeast cell an inorganic calcium phosphate or silica shell by layer-by-layer method, and further studied the implications of this modification. The silica nanocoat could improve the thermostability and water retention capability upon drying condition.
Chapter3. A eggshell-like calcium phopahte shell was introduced on Japanese encephalitis virus by in situ biomineralization. This eggshell-like exterior significantly improved the thermostability of virus vaccine without impairing its biological characteristics. This study provides a novel approach to thermostable vaccine development in developing countries.
Chapter4. Using the human enterovirus type71(EV71) vaccine strain as a model, we suggest a combined, rational design approach to improve the thermostability and immunogenicity of live vaccines by self-biomineralization. The biomimetic peptides are rationally integrated onto the capsid of EV71by reverse genetics so that calcium phosphate mineralization can be biologically induced onto vaccine surfaces under physiological conditions, generating a mineral exterior. This engineered self-biomineralized virus was characterized in detail for its unique structural, virological and chemical properties. Analogous to many exteriors, the mineral coating confers some new properties on enclosed vaccines. Such a combination of genetic technology and biomineralization provides an economic solution for current vaccination programs, especially in developing countries that lack expensive refrigeration infrastructures.
Chapter5. We showed a feasible strategy to produce thermostable liquid vaccines by introducing a protective hydrated amorphous silica exteriors to them with tunable in situ silicification. The hydrated amorphous silica exterior does not impair the original biological behaviors of virus vaccines, but confers a significantly improved thermostability on them at a wide range of temperatures. After silicification, the storage period of Poliovirus and EV71virus at room temperature can be extended by about7-fold and10-fold respectively, realizing the storage of these vaccines at room temperature for more than one month. Silica exteriors confined nearby water molecules through hydrogen bonds formed between water and hydroxyl groups, and therefore decreased the exchange of ionic and hydrogen bonding between solution and interior vaccine. The technology may pose great impact on current vaccination program by providing an economic strategy for producing liquid thermostable vaccines. Based on these results, we suggest a new concept that the silica achors can improve organisms'thermostability. The study also provides a reasonable explanation why the nature organisms often choose amorphous silica as the biominerals.
Chapter6. We briefly summaried the studies in this dissertation. Our studies clearly demostrate that the biomimetic mineralization strategy can be successfully applied in improving organisms' themostability, and hold great promise in the development of liquid thermostable vaccines. We further disscussed how to better applied this functional materials based biomimetic strategy to improve the biologicals' applications in future.
引文
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