Lesson Two
Photosynthesis
内容：
Photosynthesis occurs only in the chlorophyllchlorophyll叶绿素-containing cells of green plants, algae藻, and certain protists 原生生物and bacteria. Overall, it is a process that converts light energy into chemical energy that is stored in the molecular bonds. From the point of view of chemistry and energetics, it is the opposite of cellular respiration. Whereas 然而 cellular细胞的 respiration 呼吸is highly exergonic吸收能量的 and releases energy, photosynthesis光合作用 requires energy and is highly endergonic.
光合作用只发生在含有叶绿素的绿植物细胞，海藻，某些原生动物和细菌之中。总体来说，这是一个将光能转化成化学能，并将能量贮存在分子键中，从化学和动能学角度来看，它是细胞呼吸作用的对立面。细胞呼吸作用是高度放能的，光合作用是需要能量并高吸能的过程。
Photosynthesis starts with CO2 and H2O as raw materials and proceeds through two sets of partial reactions. In the first set, called the light-dependent reactions, water molecules are split裂开 (oxidized), 02 is released, and ATP and NADPH are formed. These reactions must take place in the presence of 在面前 light energy. In the second set, called light-independent reactions, CO2 is reduced (via the addition of H atoms) to carbohydrate. These chemical events rely on the electron carrier NADPH and ATP generated by the first set of reactions.
光合作用以二氧化碳和水为原材料并经历两步化学反应。第一步，称光反应，水分子分解，氧分子释放，ATP和NADPH形成。此反应需要光能的存在。第二步，称暗反应，二氧化碳被还原成碳水化合物，这步反应依赖电子载体NADPH以及第一步反应产生的ATP。
Both sets of reactions take place in chloroplasts. Most of the enzymes and pigments 素for the lightdependent reactions are embedded 深入的 内含的in the thylakoid类囊体 membrane膜 隔膜 of chloroplasts 叶绿体. The dark reactions take place in the stroma.基质
两步反应都发生在叶绿体中。光反应需要的大部分酶和素包埋在叶绿体的类囊体膜上。暗反应发生在基质中。
光合作用以二氧化碳和水为原材料并经历两步化学反应。第一步，称光反应，水分子分解，氧分子释放，ATP和NADPH形成。此反应需要光能的存在。第二步，称暗反应，二氧化碳被还原成碳水化合物，这步反应依赖电子载体NADPH以及第一步反应产生的ATP。
Both sets of reactions take place in chloroplasts. Most of the enzymes and pigments 素for the lightdependent reactions are embedded 深入的 内含的in the thylakoid类囊体 membrane膜 隔膜 of chloroplasts 叶绿体. The dark reactions take place in the stroma.基质
两步反应都发生在叶绿体中。光反应需要的大部分酶和素包埋在叶绿体的类囊体膜上。暗反应发生在基质中。
How Light Energy Reaches Photosynthetic Cells（光合细胞如何吸收光能的）
The energy in light photons in the visible part of the spectrum can be captured by biological molecules to do constructive work. The pigment chlorophyll in plant cells absorbs photons within a particular absorption spectrums statement of the amount of light absorbed by chlorophyll at different wavelengths. When light is absorbed it alters the arrangement of electrons in the absorbing molecule. The added energy of the photon boosts the energy condition of the molecule from a stable state to a less-stable excited state. During the light-dependent reactions of photosynthesis, as the absorbing molecule returns to the ground state, the "excess" excitation energy is transmitted to other molecules and stored as chemical energy.
生物分子能捕获可见光谱中的光能。植物细胞中叶绿素在不同光波下吸收部分吸收光谱。在吸收分子中，光的作用使分子中的电子发生重排。光子的能量激活了分子的能量状态，使其从稳定态进入不稳定的激活态。
All photosynthetic organisms contain various classes of chlorophylls and one or more carotenoid (accessory) pigments that also contribute to photosynthesis. Groups of pigment
The energy in light photons in the visible part of the spectrum can be captured by biological molecules to do constructive work. The pigment chlorophyll in plant cells absorbs photons within a particular absorption spectrums statement of the amount of light absorbed by chlorophyll at different wavelengths. When light is absorbed it alters the arrangement of electrons in the absorbing molecule. The added energy of the photon boosts the energy condition of the molecule from a stable state to a less-stable excited state. During the light-dependent reactions of photosynthesis, as the absorbing molecule returns to the ground state, the "excess" excitation energy is transmitted to other molecules and stored as chemical energy.
生物分子能捕获可见光谱中的光能。植物细胞中叶绿素在不同光波下吸收部分吸收光谱。在吸收分子中，光的作用使分子中的电子发生重排。光子的能量激活了分子的能量状态，使其从稳定态进入不稳定的激活态。
All photosynthetic organisms contain various classes of chlorophylls and one or more carotenoid (accessory) pigments that also contribute to photosynthesis. Groups of pigment
molecules called antenna complexes are present on thylakoids. Light striking any one of the pigment molecules is funneled to a special chlorophyll a molecule, termed a reaction-center chlorophyll, which directly participates in photosynthesis. Most photosynthetic organisms possess two types of reaction-center chlorophylls, P680 and P700, each associated with an electron acceptor molecule and an electron donor. These aggregations are known respectively as photosystem Ⅰ (P700) and photosystem Ⅱ (P680).
所有的光合作用生物含有不同等级的叶绿素和一个或多个类胡萝卜素（光合作用的辅助素）。称作天线复合体的素分子存在于类囊体中。激活素分子的光能进入叶绿素反应中心，其直接参与光合作用。大部分光反应细胞器拥有两套反应中心，P680和P700，每个光系统都含有一个电子受体和电子供体。这些集合体就是大家熟识的光合系统Ⅰ和光合系统reacttomotion翻译Ⅱ 。
The Light-Dependent Reaction: Converting Solar Energy into Chemical-Bond Energy
光反应：光能转化成化学键能
The photosystems of the light-dependent reactions are responsible for the packaging of light energy in the chemical compounds ATP and NADPH. This packaging takes place thro
所有的光合作用生物含有不同等级的叶绿素和一个或多个类胡萝卜素（光合作用的辅助素）。称作天线复合体的素分子存在于类囊体中。激活素分子的光能进入叶绿素反应中心，其直接参与光合作用。大部分光反应细胞器拥有两套反应中心，P680和P700，每个光系统都含有一个电子受体和电子供体。这些集合体就是大家熟识的光合系统Ⅰ和光合系统reacttomotion翻译Ⅱ 。
The Light-Dependent Reaction: Converting Solar Energy into Chemical-Bond Energy
光反应：光能转化成化学键能
The photosystems of the light-dependent reactions are responsible for the packaging of light energy in the chemical compounds ATP and NADPH. This packaging takes place thro
ugh a series of oxidation reduction reactions set in motion when light strikes the P680 reaction center in photosystem Ⅱ. In this initial event water molecules are cleaved, oxygen is released, and electrons are donated. These electrons are accepted first by plastoquinone and then by a series of carriers as they descend an electron transport chain. For each four electrons that pass down the chain, two ATPs are formed. The last acceptor in the chain is the P700 reaction center of photosystem Ⅰ. At this point incoming photons boost the energy of the electrons, and they are accepted by ferredoxin. Ferredoxin is then reoxidized, and the coenzyme NADP+ is reduced to the NADPH. The ATP generated previously and the NADPH then take part in the light independent reactions.
光反应的光系统将光能转化成化学复合物ATP和NADPH。当光激活光系统Ⅱ的光反应中心时，通过一系列的氧化还原反应实现能量的传递。反应开始时，水被分解，氧被释放并提供电子。电子首先传递给质体醌，然后通过一系列载体形成的电子传递链。每传递4个电子，形成2个ATP。最后一个受体存在于光反应系统Ⅰ的反应中心里。此处光子激活电子，电子传递给铁氧还蛋白。铁氧还蛋白再氧化，并且辅酶NADP+还原成NADPH。早期产生的ATP和NADPH进入暗反应。
光反应的光系统将光能转化成化学复合物ATP和NADPH。当光激活光系统Ⅱ的光反应中心时，通过一系列的氧化还原反应实现能量的传递。反应开始时，水被分解，氧被释放并提供电子。电子首先传递给质体醌，然后通过一系列载体形成的电子传递链。每传递4个电子，形成2个ATP。最后一个受体存在于光反应系统Ⅰ的反应中心里。此处光子激活电子，电子传递给铁氧还蛋白。铁氧还蛋白再氧化，并且辅酶NADP+还原成NADPH。早期产生的ATP和NADPH进入暗反应。
The production of ATP from the transport of electrons excited by light energy down an electron transport chain is termed photophosphorylation. The one-way flow of electrons through photosystems II and I is called noncyclic photophosphorylation; plants also derive additional ATP through cyclic photophosphorylation, in which some electrons are shunted back through the electron transport chain between photosystems Ⅱ and Ⅰ.
由电子传递链偶连产生ATP的过程称为光合磷酸化。通过光合系统Ⅱ流经光合系统Ⅰ的电子路径称非循环式光合磷酸化；植物通过循环式光合磷酸化获得额外的ATP，一些电子在光合系统Ⅰ和Ⅱ之间的电子传递链中回流。
The Light-Independent Reactions: Building Carbohydrates
暗反应：碳水化合物的形成
In the light-independent reactions of photosynthesis, which are driven by ATP and NADPH, C02 is converted to carbohydrate. The reactions are also known as the Calvin-Benson cycle. Atmospheric CO2, is fixed as it reacts with ribulose biphosphate (RuBP), a reaction that is catalyzed by the enzyme ribulose biphosphate carboxylase. The reduction Of C02 to carbohydrate (fructose diphosphate) is completed via several more steps of the cycle. Final
由电子传递链偶连产生ATP的过程称为光合磷酸化。通过光合系统Ⅱ流经光合系统Ⅰ的电子路径称非循环式光合磷酸化；植物通过循环式光合磷酸化获得额外的ATP，一些电子在光合系统Ⅰ和Ⅱ之间的电子传递链中回流。
The Light-Independent Reactions: Building Carbohydrates
暗反应：碳水化合物的形成
In the light-independent reactions of photosynthesis, which are driven by ATP and NADPH, C02 is converted to carbohydrate. The reactions are also known as the Calvin-Benson cycle. Atmospheric CO2, is fixed as it reacts with ribulose biphosphate (RuBP), a reaction that is catalyzed by the enzyme ribulose biphosphate carboxylase. The reduction Of C02 to carbohydrate (fructose diphosphate) is completed via several more steps of the cycle. Final
ly, RUBP is regenerated so that the cycle may continue.
由ATP和NADPH驱动的暗反应中，二氧化碳转化成碳水化合物。即卡尔文循环。二磷酸核酮糖固定二氧化碳，由二磷酸核酮糖羧化酶催化。
Oxygen: An Inhibitor of photosynthesis（氧：光合作用的抑制因子）
High levels of oxygen in plant cells can disrupt photosynthesis and can also cause photorespiration-an inefficient fun of the dark reactions in which 02 is fixed rather than C02 and no carbohydrate is produced.
Reprieve from Photorespiration: The C4 Pathway
Most plants are C3 plants; they experience decreased carbohydrate production under hot, dry conditions as a result of the effects of photorespiration. Among C4 plants, however, special leaf anatomy and a unique biochemical pathway enable the plant to thrive in and conditions. Thus C4 plants lessen photorespiration by carrying out photosynthesis only in cells that are insulated from high levels of CO2. They also possess a novel mechanism for carbon fixation.
大部分植物是碳3植物，在高温干旱条件下，由于光呼吸作用而使碳水化合物的合成降低。
由ATP和NADPH驱动的暗反应中，二氧化碳转化成碳水化合物。即卡尔文循环。二磷酸核酮糖固定二氧化碳，由二磷酸核酮糖羧化酶催化。
Oxygen: An Inhibitor of photosynthesis（氧：光合作用的抑制因子）
High levels of oxygen in plant cells can disrupt photosynthesis and can also cause photorespiration-an inefficient fun of the dark reactions in which 02 is fixed rather than C02 and no carbohydrate is produced.
Reprieve from Photorespiration: The C4 Pathway
Most plants are C3 plants; they experience decreased carbohydrate production under hot, dry conditions as a result of the effects of photorespiration. Among C4 plants, however, special leaf anatomy and a unique biochemical pathway enable the plant to thrive in and conditions. Thus C4 plants lessen photorespiration by carrying out photosynthesis only in cells that are insulated from high levels of CO2. They also possess a novel mechanism for carbon fixation.
大部分植物是碳3植物，在高温干旱条件下，由于光呼吸作用而使碳水化合物的合成降低。
而在大多数的碳4植物中，由于叶脉的特殊构造和独特的化学路径使植物依然很茂盛。这是碳固定的一个新机制。
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