大型海藻细胞光生物反应器培养重要工程参数的研究
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摘要
大型海藻是海洋里的原始植物,其体内含多种重要或具潜在药用价值的初级和次生代谢产物。陆生植物稀有的次生代谢产物在藻体内含量极低,化学结构相当复杂,使直接提取或者人工合成极为困难。光生物反应器所提供的可调控和工程优化的培养环境有望成为优化次生代谢物生物合成的有效手段。大型海藻细胞光生物反应器培养是较新的研究领域,受限于生物学研究手段,目前尚未分离得到大型海藻离体培养细胞,其体外培养载体仅限于配子体细胞和微繁殖体,并且前者形态更近似细胞。
本文借助大型褐藻海带配子体细胞来模拟大型海藻细胞的生长,全面研究了光生物反应器培养涉及的各种培养条件,包括温度、光照强度、光周期、和通气方式,对藻体细胞生长和生理状态的影响;进一步在上述各适宜条件下,研究了氮磷营养盐的不同起始浓度和不同脉冲补料方式对生物量生产的影响;同时研究了细胞生长的不适环境,即搅拌曝气培养下,细胞在不同搅拌强度和搅拌时间下的响应情况,和相应的工程学和生物学解释。
本文的创新点在于系统全面研究了大型海藻体外培养载体之一的配子体细胞,在光生物反应器中培养的各种培养条件;首次系统研究了不同保种培养体系对大型海藻细胞生长和生理状态的影响;首次将无泡硅橡胶膜运用到大型海藻细胞培养中;由于藻体细胞对氮磷物质有明显的储存倾向,本文合理巧妙运用脉冲补料方式快速扩增大型海藻体外繁殖系;此外,针对大型海藻体外繁殖系在不同搅拌剪切强度和剪切时间下的响应,其系统性研究也是本文的创新点。本实验的意义在于其中各项研究方法和实验结果是未来大规模培养大型海藻细胞收获生物量和次生代谢产物的重要基础。
细胞的光合速率、色素增幅、损伤率、生长曲线、细胞形态、系统的气液传质系数等指标表明,本实验中海带配子体细胞生长的适宜温度和光周期为13℃和16 : 8 LD,其饱和光强为28μE m-2 s-1,表面曝气中的搅拌培养和摇床培养,以及鼓泡培养,系统的kLa和速率间均符合Y=aXb的多项式,在悬浮性较低(液体湍动程度不高)、气液传质速率适中的团聚状态下,海带配子体细胞生长状态良好并能获得高产生物量,如本实验膜供气培养下,生物量倍数较静止培养、鼓泡培养、搅拌培养,其增幅分别为103%、29%、69%;但低湍动条件下细胞快速生长易导致细胞聚集体过大,使其内部细胞受光照、营养和气体供应的限制,而低剪切环境可控制细胞聚集体大小,进而确保培养物持续稳定增殖,本实验验证了自行设计的改良型搅拌鼓泡式光生物反应器适宜培养海带配子体细胞。
在该生物反应器中,随着APSW人工海水中起始硝酸盐和磷酸盐浓度的升高,海带配子体细胞的比生长速率渐增,延滞期渐短,但当其浓度超过1.189 mmol N L-1和0.0742 mmol P L-1后,生物量倍数渐少,高浓度氮磷的毒性渐显,故本实验培养液内最适氮磷浓度为0.594 mmol N L-1和0.0371 mmol P L-1,相应的生物量倍数为7.2;脉冲补料培养方式下,频繁少量补加氮磷有利于氮磷吸收间的同步性,补料起点过早或过迟都会破坏这种协同性,或使吸收的氮磷作储存之用,或严重影响细胞对氮磷的吸收,当细胞生长至对数中期开始频繁补加少量氮磷,即本实验中维持培养液内氮磷浓度在其起始值的1/3至1/2之间,对于细胞增殖非常有效,和批次培养相比,其生物量增幅高达76%。
搅拌曝气式光生物反应器中60 h短期连续剪切海带配子体细胞的研究表明,培养物叶绿素含量、培养液内磷酸盐、磷酸盐浓度和细胞损伤率适宜作为细胞受剪切损伤程度的良好指标,适度的剪切利于生物量的积累和营养盐的吸收,过大剪切强度或过长剪切时间均会造成细胞生物量负增长、胞内氮池、磷池释放、细胞损伤率升高、细胞显微形态巨变等负面影响;此外,流体力学参数,即Kolmogoroff涡流尺寸、Kolmogoroff涡流速率和最大湍动力均可对这些差异作出较好的工程学解释;21 d长期连续剪切的研究表明,搅拌速率为90 rpm,即搅拌子线速率0.254 m s-1时,细胞生长状态最佳,120-270 rpm下细胞生长呈现先升后降再回升的三段态势,相应的细胞损伤率也先增后减,因为细胞壁含量在此期间先降后升或超出对照组,细胞壁含量的变化直接解释了细胞受到剪切损伤并逐步耐受剪切环境,细胞壁含量回升力度可反映细胞耐受剪切环境的程度。
Macroalgae, the primitive oceanic plants, contain numerous primary and secondary metabolites with important or potential pharmaceutical values. The extremely low content in the algae,and the complicated molecular structure of those rare secondary metabolic compounds, however, discourage the direct extraction and chemical synthesis. Photolithotrophic cultivation of macroalgal cells is an effective way to optimize culture conditions, especially those that are specifically important, in the stimulation of secondary metabolic biosynthesis, and to achieve future commercial production of relevant biochemicals. Till now, cultivation of macroalgal cells in photobioreactors is a great challenge. And because of the weak biological techniques, there are no macroalgal cells cultivated in vitro. Only gametophyte cells and microplantlets are chosen for macroalgal tissue or cell culture. And the former resemble much the real macroalgal cells.
In this study, gametophyte cells of brown macroalgae Laminaria japonica were employed to stimulate macroalgal cell growth. Effects of all conditions involved in photobioreactor cultivation, such as temperature, light intensity, photoperiod, and aeration modes, on algal cell growth rates and cell physiological status, were comprehensively studied. Further research included the effects of different initial nutrient concentration and different pulse fed-batch modes on biomass production under the above optimal cultivation conditions. In addition, the influence of agitation intensity and agitation time on cell growth and cell injury was studied. Corresponding engineering and biology explanation was specifically elucidated.
The innovation of this experiment is the comprehensive and systemic study on the key conditions of photobioreactor cultivation of the gametophyte cells. It is the first systemic study on cell growth rates and cell physiological status under different subcultural systems of macroalgal cells. Bubble-less silicone tubular membrane aeration mode for macroalgal cell culture is applied for the first time in this paper. Because algae will accumulate much more of nutrient than is required for normal metabolism, pulse fed-batch modes are felicitously applied in this research for macroalgal cell proliferation. Moreover, the systemic study on macroalgal clones’responses to different agitation intensity and the exposure time is another innovation of this experiment. This work has provided rational methods and important data for large-scale production of biomass and important secondary metabolites of macroalgal cells in the future.
Photosynthetic rate, increment of chlorophyll a, cell injury ratio, cell growth curve, cell morphology, gas-liquid mass transfer co-efficient (kLa) indicated that the optimal temperature and photoperiod of Laminaria japonica gametophytic cells were 13℃and 16:8 LD, with the light saturation point of 28μE m-2 s-1. kLa of spinning, shaking and bubbling aeration modes versus input rates accorded well with the equation of Y=aXb. High biomass production with fine macroalgal cell physiological status was obtained under the environment of low suspension (low turbulence) and moderate gas-liquid transfer rates. In this experiment, compared with static, bubbling, and spinning mode, increase of biomass accumulation under membrane aeration mode was 103%, 29%, and 69%. However, larger cell aggregates formed because of quick cell growth under lower turbulence condition. The internal cells of the large aggregates were in lack of light, nutrient and gas delivery. Low shearing, on the other hand, could contribute to moderate aggregate size, and further the assurance of continuous cell proliferation. The self-designed modified stirred tank photobioreactors (with bubbling) were proved feasible for the cultivation of Laminaria japonica gametophyte cells.
With the increase of initial nutrient concentration in the artificial Pacific seawater (APSW) medium, specific growth rate of gametophyt cells cultivated in the above photobioreactors increased gradually, its lag phase shortened step by step. When initial nutrient concentration exceeded 1.189 mmol N L-1 and 0.0742 mmol P L-1, biomass increasing times decreased, with the potential toxicity of high nutrient concentration emerged. The optimal nutrient concentration in this work was 0.594 mmol N L-1 and 0.0371 mmol P L-1,under which biomass increasing 7.2 times. As to the pulse fed-batch modes, the results showed that feeding the culture frequently with small nutrient quantity was beneficial for the synchronization between nitrate and phosphate absorption. Feeding when ambient nutrient was abundant or depleted would result in the divergence absorption between nitrante and phosphate, nutrient storage, or the decrease of nutrient absorpsion. Feeding nutrient frequently with small quantity from mid-exponential growth of macroalgal cells, that is maintaining medium nutrient concentration between 1/3 and 1/2 of its initial concentration in this study, was the most effective way for biomass production, with 76% increase of biomass accumulation compared with the batch mode.
Results of short-term (60 h) continuous agitation on Laminaria japonica gametophytic cells under stirred tank photobioreactors (with surface aeration) indicated that chlorophyll content, medium nitrate and phosphate concentration, and cell injury ratio are suitable indicators for the evaluation of cell injuries under hydrodynamic agitation shear stress. Moderate shearing was beneficial for biomass accumulation and nutrient absorption. Greater shear stress or longer shearing time would lead to negative cell growth, the release of cellular N pool and P pool, the rise of cell injury ratio, and tremendous alteration of cell micro-morphology. In addition, hydrodynamic characteristics, such as Kolmogoroff eddy sizes, Kolmogoroff eddy velocity, and maximal turbulent stress were preferable engineering explanation for those differences under different agitation speeds. Results of long-term (21 d) continuous agitation on gametophytic cells showed that larger biomass accumulation and better cell physiological status were obtained under the agitation speed of 90 rpm (the stirrer tip speed of 0.254 m s-1). A three-phase of ascending, decending, and re-ascengding trend of cell growth was presented under the agitation speeds of 120-270 rpm. Corresponding cell injury ratio increased firstly, and then decreased. These results were due to the variation of cell wall fraction (CWF) during the agitation period. CWF under the agitation speeds of 120-270 rpm decreased, and then increased, even exceeded CWF under the control. These explained well cell injury and cell resistance to hydrodynamic shear stress.
本文借助大型褐藻海带配子体细胞来模拟大型海藻细胞的生长,全面研究了光生物反应器培养涉及的各种培养条件,包括温度、光照强度、光周期、和通气方式,对藻体细胞生长和生理状态的影响;进一步在上述各适宜条件下,研究了氮磷营养盐的不同起始浓度和不同脉冲补料方式对生物量生产的影响;同时研究了细胞生长的不适环境,即搅拌曝气培养下,细胞在不同搅拌强度和搅拌时间下的响应情况,和相应的工程学和生物学解释。
本文的创新点在于系统全面研究了大型海藻体外培养载体之一的配子体细胞,在光生物反应器中培养的各种培养条件;首次系统研究了不同保种培养体系对大型海藻细胞生长和生理状态的影响;首次将无泡硅橡胶膜运用到大型海藻细胞培养中;由于藻体细胞对氮磷物质有明显的储存倾向,本文合理巧妙运用脉冲补料方式快速扩增大型海藻体外繁殖系;此外,针对大型海藻体外繁殖系在不同搅拌剪切强度和剪切时间下的响应,其系统性研究也是本文的创新点。本实验的意义在于其中各项研究方法和实验结果是未来大规模培养大型海藻细胞收获生物量和次生代谢产物的重要基础。
细胞的光合速率、色素增幅、损伤率、生长曲线、细胞形态、系统的气液传质系数等指标表明,本实验中海带配子体细胞生长的适宜温度和光周期为13℃和16 : 8 LD,其饱和光强为28μE m-2 s-1,表面曝气中的搅拌培养和摇床培养,以及鼓泡培养,系统的kLa和速率间均符合Y=aXb的多项式,在悬浮性较低(液体湍动程度不高)、气液传质速率适中的团聚状态下,海带配子体细胞生长状态良好并能获得高产生物量,如本实验膜供气培养下,生物量倍数较静止培养、鼓泡培养、搅拌培养,其增幅分别为103%、29%、69%;但低湍动条件下细胞快速生长易导致细胞聚集体过大,使其内部细胞受光照、营养和气体供应的限制,而低剪切环境可控制细胞聚集体大小,进而确保培养物持续稳定增殖,本实验验证了自行设计的改良型搅拌鼓泡式光生物反应器适宜培养海带配子体细胞。
在该生物反应器中,随着APSW人工海水中起始硝酸盐和磷酸盐浓度的升高,海带配子体细胞的比生长速率渐增,延滞期渐短,但当其浓度超过1.189 mmol N L-1和0.0742 mmol P L-1后,生物量倍数渐少,高浓度氮磷的毒性渐显,故本实验培养液内最适氮磷浓度为0.594 mmol N L-1和0.0371 mmol P L-1,相应的生物量倍数为7.2;脉冲补料培养方式下,频繁少量补加氮磷有利于氮磷吸收间的同步性,补料起点过早或过迟都会破坏这种协同性,或使吸收的氮磷作储存之用,或严重影响细胞对氮磷的吸收,当细胞生长至对数中期开始频繁补加少量氮磷,即本实验中维持培养液内氮磷浓度在其起始值的1/3至1/2之间,对于细胞增殖非常有效,和批次培养相比,其生物量增幅高达76%。
搅拌曝气式光生物反应器中60 h短期连续剪切海带配子体细胞的研究表明,培养物叶绿素含量、培养液内磷酸盐、磷酸盐浓度和细胞损伤率适宜作为细胞受剪切损伤程度的良好指标,适度的剪切利于生物量的积累和营养盐的吸收,过大剪切强度或过长剪切时间均会造成细胞生物量负增长、胞内氮池、磷池释放、细胞损伤率升高、细胞显微形态巨变等负面影响;此外,流体力学参数,即Kolmogoroff涡流尺寸、Kolmogoroff涡流速率和最大湍动力均可对这些差异作出较好的工程学解释;21 d长期连续剪切的研究表明,搅拌速率为90 rpm,即搅拌子线速率0.254 m s-1时,细胞生长状态最佳,120-270 rpm下细胞生长呈现先升后降再回升的三段态势,相应的细胞损伤率也先增后减,因为细胞壁含量在此期间先降后升或超出对照组,细胞壁含量的变化直接解释了细胞受到剪切损伤并逐步耐受剪切环境,细胞壁含量回升力度可反映细胞耐受剪切环境的程度。
Macroalgae, the primitive oceanic plants, contain numerous primary and secondary metabolites with important or potential pharmaceutical values. The extremely low content in the algae,and the complicated molecular structure of those rare secondary metabolic compounds, however, discourage the direct extraction and chemical synthesis. Photolithotrophic cultivation of macroalgal cells is an effective way to optimize culture conditions, especially those that are specifically important, in the stimulation of secondary metabolic biosynthesis, and to achieve future commercial production of relevant biochemicals. Till now, cultivation of macroalgal cells in photobioreactors is a great challenge. And because of the weak biological techniques, there are no macroalgal cells cultivated in vitro. Only gametophyte cells and microplantlets are chosen for macroalgal tissue or cell culture. And the former resemble much the real macroalgal cells.
In this study, gametophyte cells of brown macroalgae Laminaria japonica were employed to stimulate macroalgal cell growth. Effects of all conditions involved in photobioreactor cultivation, such as temperature, light intensity, photoperiod, and aeration modes, on algal cell growth rates and cell physiological status, were comprehensively studied. Further research included the effects of different initial nutrient concentration and different pulse fed-batch modes on biomass production under the above optimal cultivation conditions. In addition, the influence of agitation intensity and agitation time on cell growth and cell injury was studied. Corresponding engineering and biology explanation was specifically elucidated.
The innovation of this experiment is the comprehensive and systemic study on the key conditions of photobioreactor cultivation of the gametophyte cells. It is the first systemic study on cell growth rates and cell physiological status under different subcultural systems of macroalgal cells. Bubble-less silicone tubular membrane aeration mode for macroalgal cell culture is applied for the first time in this paper. Because algae will accumulate much more of nutrient than is required for normal metabolism, pulse fed-batch modes are felicitously applied in this research for macroalgal cell proliferation. Moreover, the systemic study on macroalgal clones’responses to different agitation intensity and the exposure time is another innovation of this experiment. This work has provided rational methods and important data for large-scale production of biomass and important secondary metabolites of macroalgal cells in the future.
Photosynthetic rate, increment of chlorophyll a, cell injury ratio, cell growth curve, cell morphology, gas-liquid mass transfer co-efficient (kLa) indicated that the optimal temperature and photoperiod of Laminaria japonica gametophytic cells were 13℃and 16:8 LD, with the light saturation point of 28μE m-2 s-1. kLa of spinning, shaking and bubbling aeration modes versus input rates accorded well with the equation of Y=aXb. High biomass production with fine macroalgal cell physiological status was obtained under the environment of low suspension (low turbulence) and moderate gas-liquid transfer rates. In this experiment, compared with static, bubbling, and spinning mode, increase of biomass accumulation under membrane aeration mode was 103%, 29%, and 69%. However, larger cell aggregates formed because of quick cell growth under lower turbulence condition. The internal cells of the large aggregates were in lack of light, nutrient and gas delivery. Low shearing, on the other hand, could contribute to moderate aggregate size, and further the assurance of continuous cell proliferation. The self-designed modified stirred tank photobioreactors (with bubbling) were proved feasible for the cultivation of Laminaria japonica gametophyte cells.
With the increase of initial nutrient concentration in the artificial Pacific seawater (APSW) medium, specific growth rate of gametophyt cells cultivated in the above photobioreactors increased gradually, its lag phase shortened step by step. When initial nutrient concentration exceeded 1.189 mmol N L-1 and 0.0742 mmol P L-1, biomass increasing times decreased, with the potential toxicity of high nutrient concentration emerged. The optimal nutrient concentration in this work was 0.594 mmol N L-1 and 0.0371 mmol P L-1,under which biomass increasing 7.2 times. As to the pulse fed-batch modes, the results showed that feeding the culture frequently with small nutrient quantity was beneficial for the synchronization between nitrate and phosphate absorption. Feeding when ambient nutrient was abundant or depleted would result in the divergence absorption between nitrante and phosphate, nutrient storage, or the decrease of nutrient absorpsion. Feeding nutrient frequently with small quantity from mid-exponential growth of macroalgal cells, that is maintaining medium nutrient concentration between 1/3 and 1/2 of its initial concentration in this study, was the most effective way for biomass production, with 76% increase of biomass accumulation compared with the batch mode.
Results of short-term (60 h) continuous agitation on Laminaria japonica gametophytic cells under stirred tank photobioreactors (with surface aeration) indicated that chlorophyll content, medium nitrate and phosphate concentration, and cell injury ratio are suitable indicators for the evaluation of cell injuries under hydrodynamic agitation shear stress. Moderate shearing was beneficial for biomass accumulation and nutrient absorption. Greater shear stress or longer shearing time would lead to negative cell growth, the release of cellular N pool and P pool, the rise of cell injury ratio, and tremendous alteration of cell micro-morphology. In addition, hydrodynamic characteristics, such as Kolmogoroff eddy sizes, Kolmogoroff eddy velocity, and maximal turbulent stress were preferable engineering explanation for those differences under different agitation speeds. Results of long-term (21 d) continuous agitation on gametophytic cells showed that larger biomass accumulation and better cell physiological status were obtained under the agitation speed of 90 rpm (the stirrer tip speed of 0.254 m s-1). A three-phase of ascending, decending, and re-ascengding trend of cell growth was presented under the agitation speeds of 120-270 rpm. Corresponding cell injury ratio increased firstly, and then decreased. These results were due to the variation of cell wall fraction (CWF) during the agitation period. CWF under the agitation speeds of 120-270 rpm decreased, and then increased, even exceeded CWF under the control. These explained well cell injury and cell resistance to hydrodynamic shear stress.
引文
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