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生物专业英语课文翻译

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2021-02-17 07:20
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2021年2月17日发(作者:swift是什么意思)


第一课



Cytoplasm: The Dynamic, Mobile Factory



细胞质:动力工厂



Most


of


the


properties


we


associate


with


life


are


properties


of


the


cytoplasm. Much


of


the


mass


of


a


cell


consists


of


this


semifluid substance, which is bounded on the outside by the plasma membrane. Organelles are suspended within it, supported


by the filamentous network of the cytoskeleton. Dissolved in the cytoplasmic fluid are nutrients, ions, soluble proteins, and


other materials needed for cell functioning.



生命的大部分 特征表现在细胞质的特征上。细胞质大部分由半流体物质组成,并由细胞膜(原生质膜)包被。细胞器

< p>
悬浮在其中,并由丝状的细胞骨架支撑。细胞质中溶解了大量的营养物质,离子,可溶蛋白以及维持 细胞生理需求的


其它物质。



The Nucleus: Information Central


(细胞核:信息中心)



The eukaryotic cell nucleus is the largest organelle and houses the genetic material (DNA) on chromosomes. (In prokaryotes


the hereditary material is found in the nucleoid.) The nucleus also contains one or two organelles-the nucleoli-that play a role in


cell


division. A


pore-perforated


sac


called


the


nuclear


envelope


separates


the


nucleus


and


its


contents


from


the


cytoplasm. Small molecules can pass through the nuclear envelope, but larger molecules such as mRNA and ribosomes must


enter and exit via the pores.



真核细胞 的细胞核是最大的细胞器,细胞核对染色体组有保护作用(原核细胞的遗传物质存在于拟核中)

< br>。细胞核含有


一或二个核仁,核仁促进细胞分裂。核膜贯穿许多小孔,小分子可以 自由通过核膜,而象


mRNA


和核糖体等大分子必


须通过核孔运输。



Organelles: Specialized Work Units


(细胞器:特殊的功能单位)



All eukaryotic cells contain most of the various kinds of organelles, and each organelle performs a specialized function in the


cell. Organelles


described


in


this


section


include


ribosomes,


the


endoplasmic


reticulum,


the


Golgi


complex,


vacuoles,


lysosomes, mitochondria, and the plastids of plant cells.



所有的真核细胞都含有多种细胞器,每个细胞器 都有其特定功能。本节主要介绍核糖体,内质网,高尔基体系,液泡,


溶酶体,线粒体和 植物细胞中的质体。



The number of ribosomes within a cell may range from a few hundred to many thousands. This quantity reflects the fact that,


ribosomes are the sites at which amino acids are assembled into proteins for export or for use in cell processes. A complete


ribosome is composed of one larger and one smaller subunit. During protein synthesis the two subunits move along a strand of


mRNA,


attached


to


a


single


mRNA


strand;


such


a


combination


is


called


a


polysome. Most


cellular


proteins


are


manufactured


on


ribosomes in the cytoplasm. Exportable proteins and membrane proteins are usually made in association with the endoplasmic


reticulum.



核糖体的数量 变化从几百到几千,核糖体是氨基酸组装成蛋白质的重要场所。完整的核糖体由大亚基和小亚基组成。

< p>
核糖体沿着


mRNA


移动并阅读遗传密码,


翻译成蛋白质。


一条


mRNA


上可能有多个核糖体,


称多聚核糖体。


大多数细


胞蛋白是由细胞质中核糖体生产。输出蛋白和膜蛋白通常与内质网有关。



The


endoplasmic


reticulum,


a


lacy


array


of


membranous


sacs,


tubules,


and


vesicles,


may


be


either


rough


(RER)


or


smooth


(SER). Both types play roles in the synthesis and transport of proteins. The RER, which is studded with polysomes, also seems


to be the source of the nuclear envelope after a cell divides.


< br>内质网,带有花边的生物囊,有管状,泡状之分,以及光滑和粗糙面区别。两种都与蛋白质的合成和运输有 关。粗糙


内质网上分布许多核糖体,也可能提供细胞分裂后所需的细胞膜。



SER lacks polysomes; it is active in the synthesis of fats and steroids and in the oxidation of toxic substances in the cell. Both


types of endoplasmic reticulum serve as compartments within the cell where specific products can be isolated and subsequently


shunted to particular areas in or outside the cell.



光滑内质网上无核糖体 ,主要作用是脂肪和类固醇的合成以及细胞内有毒物质的氧化。两种内质网合成的产物在其中

进行分流或运输到细胞外。



Transport vesicles may carry exportable molecules from the endoplasmic reticulum to another membranous organelle, the Golgi


complex. Within the Golgi complex molecules are modified and packaged for export out of the cell or for delivery else where


in the cytoplasm.



运输小泡能够将可运输分子从内质网运 输到高尔基复合体上。在高尔基复合体中修饰,包装后输出细胞或传递到细胞


1


/


20


质中的其他场所。



Vacuoles


in


cells


appear


to


be


hollow


sacs


but


are


actually


filled


with


fluid


and


soluble


molecules. The


most


prominent


vacuoles


appear


in


plant


cells


and


serve


as


water


reservoirs


and


storage


sites


for


sugars


and


other


molecules. Vacuoles


in


animal cells carry out phagocytosis (the intake of particulate matter) and pinocytosis (vacuolar drinking).



细胞中的液泡好象是中空的,但实际上充满了液体和可溶分子。最典型的液泡存在于植物细胞 中,储备水,糖以及其


它分子。动物中的液泡起吞噬和胞饮作用。



A subset of vacuoles are the organelles known as lysosomes, which contain digestive enzymes (packaged in lysosomes in the


Golgi


complex)


that


can


break


down


most


biological


macromolecules. They


act


to


digest


food


particles


and


to


degrade


damaged cell parts.



溶酶体是液泡亚单位,含有消化酶,降解大部分 生物大分子。消化食物微粒和降解损伤的细胞残片。



Mitochondria are the sites of energy- yielding chemical reactions in all cells. In addition, plant cells contain plastids that utilize


light energy to manufacture carbohydrates in the process of photosynthesis. It is on the large surface area provided by the inner


cristae of mitochondria that ATP- generating enzymes are located. Mitochondria are self-replicating, and probably they are the


evolutionary descendants of what were once free-living prokaryotes.


线粒体是细胞中化学产能的场所。另外,植物细胞中的质体在光合作用中利用光能产生碳水化合物,线粒体内 嵴上提


供了很大的表面积并分布着产


ATP

酶。线粒体自我复制,并且可能是自由生活的原核生物在进化中形成的后代。



There are two types of plastids: leucoplasts, which lack pigments and serve as storage sites for starch, proteins, and oils; and


chromoplasts,


which


contain


pigments. The


most


important


chromoplasts


are


chloroplasts-organelles


that


contain


the


chlorophyll used in photosynthesis. The internal structure of chloroplasts includes stacks of membranes called grana, which are


embedded in a matrix called the stroma.



质体有两种类型:白色体,缺乏色素,是淀粉 ,蛋白质和油的储备场所;色质体,含有色素。叶绿体是最重要的色质


体,含有与光合作 用有关的叶绿素。叶绿体的内部结构是由多层膜形成的叶绿体基粒,其中包埋在基质中的基粒称子


座。



The Cytoskeleton


(细胞骨架)



All


eukaryotic


cells


have


a


cytoskeleton,


which


is


a


convoluted


latticework


of


filaments


and


tubules


that


appears


to


fill


all


available space in the cell and provides support for various other organelles. A large portion of the cytoskeleton consists of


threadlike microfilaments composed mainly of the contractile protein actin. They are involved in many types of intracellular


movements in plant and animal cells. A second protein, myosin, is involve in the contraction of muscle cells. Another main


structural


component


of


the


cytoskeleton


consists


of


microtubules,


which


are


composed


of


the


globular


protein


tubulin


and


together


act


as


scaffolding


that


provides


a


stable


cell


shape. Cytoskeletal


intermediate


filaments


appear


to


impart


tensile


strength to the cell cytoplasm. Mechanoenzymes such as myosin, dynein, and kinesin interact with the cytoskeletal filaments


and tubules to generate forces that cause movements.



所有的细胞都有细胞骨架,网络结构的纤丝充满了它所能触及的全部空间并且对细胞器提供支持作用。细胞骨架 大部


分由微丝组成,微丝主要由可收缩的肌动蛋白组成。动植物细胞的许多种类型细胞内 运动与肌动蛋白有关。第二类蛋


白是肌球蛋白,它与肌肉细胞的收缩有关。细胞骨架的另 一个主要结构成分是微管,由球状的微管蛋白组成,象脚手


架一般维持细胞的稳定形态。 细胞骨架的中间丝提供了细胞质伸缩动力。机械酶,例如,肌球蛋白,动力蛋白,驱动


蛋 白与微丝,微管相互作用产生动力而引起细胞运动。



Cellular Movements


(细胞运动)



Although the cytoskeleton provides some stability to cells, its microtubules and filaments and their associated proteins enable


cells to move by creeping or gliding. Such movements require a solid substrate to which the cell can adhere and can be guided


by the geometry of the surface. Some cells also exhibit chemotaxis, the ability to move toward or away from the source of a


diffusing chemical.


< br>尽管细胞骨架提供了细胞的某些稳定性,微丝,微管及相关蛋白能使细胞爬行或滑动。这种运动需要固体基 质依托并


通过表面几何形状的改变而运动。某些细胞具备趋药性,即趋向或逃离扩散开的 化学源。



Certain eukaryotic cells can swim freely in liquid environments, propelled by whiplike cilia or flagella. Both cilia and flagella


have the same internal structure: nine doublets (pairs of microtubules) are arranged in a ring and extend the length of the cilium


or flagellum, and two more microtubules run down the center of the ring. Every cilium or flagellum grows only from the cell


surface where a basal body is located. Movement is based on the activities of tiny dynein side arms that extend from one of the


microtubules of each doublet.



2


/


20


某些真核细胞能在液体液体中自由运动,由纤毛或鞭毛推 动。纤毛和鞭毛具有同样的内部结构:九个双微管环形排列,


纵向延伸,环中心是两个或 以上微管组成。纤毛或鞭毛从细胞表面的基体出生长,双微管的动力蛋白臂从一侧延伸到


另一侧而引起运动。



Nutrients, proteins, and other materials within


most


plant cells are


moved about via cytoplasmic streaming. The process


occurs as myosin proteins attached to organelles push against microfilaments arrayed throughout the cell. Microfilaments and


microtubules are responsible for almost all


major cytoplasmic


movements. During cell division,


microtubules


of the spindle


assembled from tubutin subunits near organelles called centrioles move the chromosomes.



大部分植物细胞的营养,蛋白质和 其它物质由细胞质流运输。这个过程是由于依附在细胞器上的肌球蛋白反推排列在


细胞周 围的微丝形成的。绝大部分细胞质运动由微丝和微管完成。在细胞分裂期间,中心粒周围的由微管蛋白亚基装


配形成的纺锤体微管移向染色体。



第二课



Photosynthesis occurs only in the chlorophyll-containing cells of green plants, algae, and certain protists 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 cellular respiration is highly exergonic and releases


energy, photosynthesis requires energy and is highly endergonic.



光合作用只发生在含有叶绿素的绿色 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


(oxidized),


02


is


released,


and


ATP


and


NADPH


are


formed. These reactions must take place in the presence of 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



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


for


the


lightdependent


reactions


are


embedded in the thylakoid membrane of chloroplasts. The dark reactions take place in the stroma.



两步反应都发生在叶绿体中。光反应需要的大部分酶和色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



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




(P700) and photosystem




(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(光合作用的辅助色素)


。称作天线复合 体的色素


分子群存在于类囊体中。激活色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< /p>



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Ⅱ。



The Light-Dependent Reaction: Converting Solar Energy into Chemical-Bond Energy



3


/


20


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 through a series of oxidation reduction reactions set in motion when light strikes


the P680 reaction center in photosystem



. 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



.


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



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

< br>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



ATP

< p>
。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,< /p>


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



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




and



.



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. Finally, RUBP is regenerated so that the


cycle may continue.


< br>由


ATP



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.


< br>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,< /p>


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。



第三课



The Nucleus and Chromosomes



The cell nucleus is the main repository of genetic information. Within the nucleus are the chromosomes tightly coiled strands


of


DNA


and


clusters


of associated


proteins. Long


stretches


of


the


continuous DNA


molecule


wind


around


these


clusters


of


proteins,


or


histones,


forming


beadlike


complexes


known


as


nucleosomes.


More


coiling


and


supercoiling


produces


a


dense


chromosome structure. Each long strand of DNA combines with histones and nonhistone proteins to make up the substance


chromatin.



细胞核是贮藏 遗传信息的主要场所。


DNA


盘绕成螺旋线以及相关的成簇蛋白 质。


DNA


螺旋线缠绕成簇的组蛋白形成


珠链状的核小体。


这些螺旋和超螺旋形成致密的染色体组结构。


每个长链


DNA


与组蛋白和非组蛋白一起构成染色质物


质。



4


/


20


A pictorial display of an organism's chromosomes in the coiled, condensed state is known as a karyotype. Karyotype reveal


that in most cells all but sex chromosomes are present as two copies, referred to as homologous pairs. Non-sex chromosomes


are


called


autosomes. Organisms


whose


cells


contain


two


sets


of


parental


chromosomes


are


called


diploid;


those


with


cells


containing a single set of parental chromosomes are called haploid.



染色体致密的超螺旋状态我们称染色体组。除了性染色体外,大多数细胞的染色体组成对出现,称同源染 色体对。非性染色


体称常染色体。生物细胞含有两套父母本染色体的称二倍体;含有单套 染色体的称单倍体。



The Cell Cycle



The cell cycle is a regular sequence in which the cell grows, prepares for division, and divides to form two daughter cells, each


of which then repeats the cycle. Such cycling in effect makes single-celled organisms immortal. Many cells in multicellular


organisms, including animal muscle and nerve cells, either slow the cycle or break out of it altogether.



在细胞生长过程中,细胞循环 遵循特定程序,分裂准备,分裂成


2


个子细胞,子细胞再循环。 此循环使得单细胞永生。


多细胞生物中的许多细胞,包括动物肌肉和神经细胞,要么降低 循环速度,要么同时分裂。



The normal cell cycle consists of four phases. The first three include G1, the period of normal metabolism; S phase, during


which normal synthesis of


biological molecules continues, DNA is replicated, and histones are synthesized; and G2, a brief


period of metabolism and additional growth. Together the G1, S, and G2 phases are called interphase. The fourth phase of the


cell


cycle


is


M


phase,


the


period


of


mitosis,


during


which


the


replicated


chromosomes


condense


and


move


and


the


cell


divides. It is believed that properties of the cell cytoplasm control the cell cycle, along with external stimulators and inhibitors


such as chalones.



正常细胞循环由


4


个时期组成。


头三期包括


G1,


正常新陈代谢;

< p>
S



,


正常新陈代谢同时 ,


DNA


复制,


组蛋白合成;



G2


期,短期的新陈代谢和少许生长。


G1,


S,



G2


称分裂间期。最后是


M


期,有丝分裂期,复制的染色体组浓缩, 移动


并细胞分裂。据称是染色质控制了细胞循环,伴随外部激活因子和抑制因子如抑素。



Mitosis: Partitioning the Hereditary Material



Biologists divide the mitotic cycle into four phases. At the beginning of prophase the chromosomes each consist of two highly


condensed


chromatids


attached


to


each


other


at


a


centromere. As


prophase


ends


and


metaphase


begins,


the


condensed


chromosomes


become


associated


with


the


spindle. Eventually


the


chromosomes


become


arranged


in


a


plane


(called


the


metaphase plate) at a right angle to the spindle fibers. Next, during anaphase, the two sister chromatids of each chromosome


split, and one from each pair is drawn toward each pole of the cell. During telophase nuclear envelopes begin to form around


each set of chromosomes, and division of the cytoplasm takes place.


生物学家将有丝分裂划分为


4


个阶段。分裂前期,高度浓缩 的两个染色单体通过着丝粒连接在一起。在分裂前期后期


和分裂中期前期,浓缩的染色体 与纺锤体相连,最后以正确的角度排列在赤道板上。在分裂后期,两个姊妹单体分离,


分 别拽向细胞两极。在分裂末期,在每套染色体周围形成核膜,细胞质发生分裂。



As mitosis proceeds, the spindle microtubules play a crucial role in ensuring that both paired and separated chromatids move in


the right directions at the proper times. Each half of the spindle forms as microtubules extend from each pole of a dividing cell


to


the


region of


the


metaphase


plate. During


prophase,


other


microtubules,


the


centromeric


fibers,


extend


outward


from


the


spindle


poles


to


structures


on


the


chromosomes


called


kinetochores. During


anaphase


the


fibers


begin


to


shorten,


and


the


chromatids begin to move apart.



在有丝分裂过程中,是纺锤体微管确保了染色单 体在适当时间以正确方向进行分离。纺锤体微管由两极向赤道板延伸。


在分裂前期,其它 微管,着丝粒纤维延伸到染色体的动粒。在分裂后期,纤维开始变短,染色单体分离。



The spindle forms differently in plant and animal cells. In animals it is associated with centriole, while in plant and fungal cells


spindle formation is associated with reions called microtubule organizing centers.



植物和动物细胞形成的纺锤体不同。动物细 胞与中心粒相连,而在植物和真菌细胞中,纺锤体与微管组织中心的离子相连。



Cytokinesis: Partitioning the Cytoplasm



胞质分裂:细胞质分离



The division of the cell cytoplasm at the end of mitosis is called cytokinesis. In animal cells it takes place as a ring of actin


filaments


contracts


around


the


cell


equator,


pinching


the


cell


in


two. In


plant


cells,


which


are


bounded


by


a


cell


wall,


cytokinesis involves the building of a new cell plate across the dividing cell at its equator. Cell wall material is then deposited


in the region of the cell plate.



在动物细胞中,环形肌动蛋白丝延赤道板收缩而 使细胞一分为二。在植物细胞中,在赤道板形成新的细胞板。



Meiosis: The Basis of Sexual Reproduction



5


/


20


Meiosis is a special form of cell division that takes place in the reproductive organs that produce sex cells. Like mitosis, it


takes


place


after


DNA


replication


has


occurred


and


involves


two


sequential


nuclear


divisions


(meiosis


I


and


meiosis



). These


divisions


result


in


four


daughter


cells,


each


with


half


the


number


of


chromosomes


of


the


parent


cell. The


phenomenon of crossing over during meiosis results in exchanges of genetic information between chromosomes. Hence, the


homologous chromosomes distributed to different progeny cells are not identical.



减数分裂是性细胞分裂的特殊形式。< /p>


如有丝分裂,


它也是发生在


DNA


复制后并有连续的两个核分裂。


产生


4


个子细胞,


分别含有亲本一半的染色体数。



As in mitosis two chromatids exist for each chromosome at the beginning of prophase 1. During this phase the homologous


chromosomes


undergo


synapsis,


or


pairing,


which


is


brought


about


by


a


bridging


structure


of


proteins


and


RNA


called


the


synaptonemal complex. The homologous pairs stay together when they align on the metaphase plate. Unlike the anaphase of


mitosis, however, during anaphase I the two chromatids of each chromosome stay joined at the centromere and move together


to one of the two poles of the cell. It is this event that results in the halving of the chromosome number in the four daughter


cells that result from meiosis.



正如在有丝分裂 中一样,两个同源染色单体通过蛋白质和


RNA


桥配对形成联会 复合体。与有丝分裂不同的是,每组染


色体的两个染色单体连接在着丝点上并一起移向细 胞两极的一级。由此而导致


4


个子细胞染色体数减半。



During


telophase


I


nuclear


envelopes


enclose


the


chromosomes


in


nuclei,


and


in


most


species


cytokinesis


(the


first


nuclear


division)


follows. The


second


nuclear


division


begins with


metaphase



,


in


which


the


chromosomes


in


each


daughter cell


again align on a metaphase plate. The centromeres finally divide, and each sister chromatid moves to one of the poles of the


spindle. The next phase is telophase , followed again by cytokinesis. The result of the entire process is four haploid cells in


which parental chromosomes are randomly distributed.



第二次核分裂开始于分裂中期,子 细胞中染色体重新排列在赤道板上。着丝粒最终分离,每个姊妹染色单体分向两极。


接着 胞质分裂。产生


4


个单倍体,父母染色体随机分配。

< p>


Asexual Versus Sexual Reproduction



Mitosis


and


meiosis,


respectively,


make


simple


cell


division


and


sexual


reproduction


possible. Each


means


of


passing


on


hereditary


information


has


advantages. In


asexual


reproduction


the


parent


organism


gives


rise


to


offspring


that


are


genetic


clones


of


the


parent. The


advantages


of


this


type


of


reproduction


are


that


it


preserves


the


parent's


successful


genetic


complement, requires little or no specialization of reproductive organs, and is more rapid than sexual reproduction. A major


disadvantage of the asexual mode is that a single catastrophic event or disease may destroy an entire population of genetically


identical organisms. A prime benefit of sexual reproduction is that it provides genetic variability and a ready mechanism for the


elimination of deleterious mutations. It also allows



有丝分裂和减数分裂在传递遗传信息过程中各有优势。体细胞的繁殖就是父母本的克隆,其优势是保留了父母本的 成功遗传


信息,不需要特殊器官,比性复制快的多。但一个简单灾难性事件或疾病都可能 摧毁一个细胞群体。性复制的优势是它提供


了遗传可变性和现存排除有害突变的机制。也 可以产生新的基因并在种群中蔓延。



第四课



Foundations of Genetics


Early Theories of inheritanee


Early ideas of inheritance included Hippocrates' theory of


pangenesis


and August Weismands


germ



plasm theory.



Based


on experiments with mice, Weismann proposed that hereditary information in gametes transmitted traits to progeny.



Both of


these early views incorporated the blending theory: they held that heritable traits of the two parents blend, so that the distinct


characteristics of each are lost in offspring.



遗传学的早期理论包括泛生说和种 质理论。基于小鼠实验,维丝曼提出遗传信息储存在配子中并将遗传信息传递给后代。


这 两个早期观点合起来形成融合理论:子代拥有父母本混合的遗传特征,而不完全象亲代。



Gregor Mendel and the Birth of Genetics


Gregor Mendel, an Augustinian monk in the monastery at Brunn, Austria, is known as the


exposed


to


theories


of


the


particulate


nature


of


matter


while


a


university


student


and


having


a


background


in


mathematics,


Mendel


carried


out


a


series


of


carefully


planned


experiments


that


demonstrated


the


particulate


nature


of


heredity.



His


revolutionary ideas were neither understood nor accepted until many years after Mendel died.



孟德尔,众所周知 的遗传学之父,是一名修道士。当他还是大学生时就提出了物质的粒子属性。孟德尔进行了一系列周密

< p>
安排的实验来证实遗传的颗粒性。直到他去世后,他的理论才被理解和接受。



6


/


20


Mendel's Classic Experiments


Mendel studied genetics through plant- breeding experiments with the garden pea, a plant species that is self-fertilizing and


breeds true (each offspring is identical to the parent in the trait of interest).



To test the blending theory, he focused his


research on seven distinct characters.



Each of these characters, such as seed color and plant height, present only two, clear- cut


possibilities.



He also recorded the type and number of all progeny produced from each pair of parent pea plants, and followed


the results of each cross for two generations.


孟德尔通过豌豆 实验研究遗传学,豌豆是自花授粉植物和纯品系。为验证融合理论,他的研究主要集中在


7


个特征上。例


如,种子颜色,植株高度,这些特征只有两个明 确的可能性。他记录了产生的每一个子代类型和数量,在杂交产生子


2

< br>代。



For each of the characters he studied, Mendel found that one trait was


dominant


while the other was


recessive.


In the second


filial


(F


2


)


generation,


the


ratio


of


dominant


to


recessive


was


3:1.


Mendel


deduced


that


this


result


was


possible


only


if


each


individual


possesses


only


two


hereditary


units,


one


from


each


parent.


The


units


Mendel


hypothesized


are


today


known


as


alleles,


alternative forms of genes.


Genes


are the basic units of heredity. An organism that inherits identical alleles for a trait


from


each


parent


is


said


to


be


homozygous


for


that


trait;


if


different


alleles


for


a


trait


are


inherited,


the


organism


is


heterozygous


for that trait. When an organism is heterozygous for a trait, the resulting phenotype for that trait expresses only


the dominant , the organism’s

phenotype



its



physical appearance and properties- differs from its


genotype,


which


may include both a dominant and a recessive allele.



A pictorial representation of all possible combinations of a genetic cross


is known as a


Punnett square.



对于每个特征而言,要么显形,要么隐性。在 子


2


代中显形与隐性比为


3

< p>


1


。只有在每个个体仅拥有两个研究遗传单元,


并每个单元来自一个亲代时,实验结果才成立。此遗传单元就是今天共识的等位基因。两 个一样的等位基因决定一个特征,


称纯合。相反,称杂合。当生物是杂合时,它的表型由 显性基因决定。因此,生物的表型与基因型是不同的。旁纳特方格可


以陈列所有可能的遗 传组合。



The


results


of


Mendel's


experiments


on


dominant


and


recessive


inheritance


let


to


Mendel's


first


law:


the


law


of


segregation.


This law states that for a given trait an organism inherits one allele from each parent.



Together these alleles form


the allele pair. When gametes are formed during meiosis, the two alleles become separated (halving of chromosome number).To


gain


evidence


for


his


theory


Mendel


performed


test


crosses,


mating


plants


of


unknown


genotype


to


plants


that


were


homozygous recessive for the trait of interest.



The ratio of dominant phenotypes (if any) in the progeny makes clear whether


the unknown genotype is heterozygous, homozygous dominant, or homozygous recessive.



分离定律,生物只遗传父母本等 位基因对的一个等位基因。减数分裂期形成配子时两个等位基因分离。为验证此理论,


他 做了测交实验,即基因型未知的植物与纯合的隐性基因植物杂交。子代显性表型可以明确测得杂合基因或纯合基因 的基因


型。



Mendel's Ideas and the Law of independent Assortment


Mendel also performed


dihybrid crosses,


which enabled him to consider how two traits are inherited relative to one another.



This work let to the


law of independent assortment,


which states that the alleles of genes governing different characters are


inherited


independently.



An


apparent


exception


to


Mendel's


laws


is


incomplete


dominance,


a


phenomenon


in


which


offspring of a cross exhibit a phenotype that is intermediate between those of the parents.



However, incomplete dominance


reflects


the


fact


that


both


alleles


for


the


trait


in


question


exert


an


effect


on


the


phenotype.



The


alleles


themselves


remain


separate.



双因子杂合试 验,两个特征是如何相互影响遗传的。试验结果产生独自分配定律,即等位基因独立遗传。特例是,不完


全显性。子代的表型是父母本的中间类型。不完全显性说明了两个等位基因对表型都有影响。


,等位基因会继续分离。



Mendel presented his ideas in 1866 in a scientific paper published by the Brunn Society for Natural History.



Unfortunately,


the meaning of his research was not understood by other scientists of the day.



His work was rediscovered in 1900 by Carl


Correns and Hugo de Vries.


1866


年,孟德尔在自然史上发表了他的 科学论文,陈诉了他的观点。不幸的是,他的研究不被当时科学家接受。在


1900


年,他的著作再被发现利用。



Chromosomes and Mendelian Genetics


Soon after Mendel's work was rediscovered, Walter Sutton and Theodor Boveri independently proposed that the hereditary


units might be located on chromosomes.



Experiments to prove this hypothesis were carried out by Thomas Hunt Morgan and


his students at Columbia University, in research on the sex chromosomes of fruit flies.



Morgan's studies were also the first


7


/


20


exploration of sex-linked traits.



It also led to the discovery in 1916 by Calvin Bridges of the phenomenon of


nondisjunction,


in which a chromosome pair fails to segregate during meiosis.


孟德尔著作被再发现不久,


Walter Sutton



Theodor Boveri


提出 ,遗传单位可能定位在染色体组上。伴性遗传又导致了


不分离现象的发现,即在减数分裂 中,染色体对不分离。



第五课



Genes Code for Particular Proteins



The


first


scientist


to


investigate


the


question


of


how


genes


affect


phenotype


was


Sir


Archibald


Garrod,


whose


studies


of


alkaptonuria implied a relationship between genes and enzymes. Thirty years later Beadle and Ephrussi showed a relationship


between


particular


genes


and


biosynthetic


reactions


responsible


for


eye


color


in


fruit


flies.


Next,


in


a


series


of


classic


experiments


on


the


effects


of


mutations


in


the


bread


mold


Neurospora


crassa,


Beadle


and


Tatum


explored


the


one- gene-one-enzyme hypothesis-the idea that each gene codes for a particular enzyme. Their work paved the way for other


researchers to elucidate the precise ways in which enzymes affect complex metabolic 1949, in research on the role


of


hemoglobin


in


sickle


cell


anemia,


Linus


Pauling


helped


refine


the


one- gene-one-enzyme


hypothesis


into


the


one- gene-one-polypeptide hypothesis.



Archibald


Garrod


是第一个研究基因是如何影响表型的科学家,他对尿黑酸症的研究揭示了基因与酶之间的关系。


Beadle



Ephrussi

< br>在三十年后对果蝇眼睛颜色的研究发现特殊基因与相关反应的生物合成有关。接着对面包发霉粗糙


脉孢菌的突变试验得出一个基因一个酶的假说。他们的工作为其他工作者铺平了道路,即精确地阐 明了酶影响了复杂


的新陈代谢途径。在


1949


年,对镰刀状细胞贫血症的研究对一个基因一个酶的假说进一步上升为一个基因一个多肽。



The Search for the Chemistry and Molecular Structure of nucleic Acids



Nuclei acid, originally isolated by Johann Miescher in 1871, was identified as a prime constituent of chromosomes through the


use


of


the


red-staining


method


developed


by


Feulgen


in


the


early


1900s. Frederick


Griffith's


experiments


with


the R


and S


stains of pneumococci showed that an as yet unknown material from one set of bacterial could alter the physical traits of a


second set. In the 1940s the team of Avery, MacLeod, and McCarty showed that this unknown material was DNA. At about


the


same


time


P.A.


Levene


discovered


that


DNA


contained


four


nitrogenous


bases,


each


of


which


was


attached


to


a


sugar


molecule and a phosphate group-a combination Levene termed a nucleotide.




18 71


年,核酸最初是由


Johann


Miescher


分离成功,并由


Feulgen



1900


年证实核酸是染色体组最基本的组成。< /p>


Frederick Griffith


对粗糙和光滑的肺炎球菌 实验表明,


不确定的某种物质可以从一组细菌转移到另一种细菌中。


1940


年,确认该物质为


DN A


。四个碱基和磷酸分子分别连接在糖分子上,称核苷酸。



Disagreement over whether DNA could carry complex genetic information was ended in the early 1950s by Martha Chase and


Alfred Hershey, whose work with E. coli showed clearly that DNA, and not protein, is the bearer of genetic information.



直到

< p>
1950


年,通过对大肠杆菌实验发现,遗传物质是


DNA


,而不是蛋白质。



Each DNA nucleotide contains a five-carbon sugar, deoxyribose, attached to one of four bases: adenine, guanine, cytosine, or


thymine. Adenine and guanine molecules are double-ring structures called purines, while cytosine and thymine are single-ring


structures called pyrimidines. The molecule made up of a base plus a sugar is termed a nucleoside. In each molecule of DNA a


phosphate group links the five-carbon sugar of one nucleoside to the five-carbon sugar of the next nucleoside in the chain. This


phosphate bonding creates a sugar-phosphate backbone.



每个核苷酸都含有一个五碳脱氧核糖,分别连接


4


个碱基,即: 腺嘌呤,鸟嘌呤,胞嘧啶,胸腺嘧啶。碱基连接糖称


核苷。磷酸键形成磷酸骨架。



Chargaff’s rules describe the fact that (1) the amount of adenine is equal to the amou


nt of thymine in DNA, with amount of


cytosine equal to that of guanine, and (2) the ratios of A to T and of C to G vary with different species.


< br>(


1


)腺嘌呤与胸腺嘧啶,胞嘧啶与鸟嘌呤相等;



2


)腺嘌呤与胸腺嘧啶,胞嘧啶与鸟嘌呤的 比例随物种不同而不同。



The Research Race for the Molecular Structure of DNA



In the late 1940s and early 1950s, researchers looking for the structure of DNA drew upon Chargaff s insight, Levene's ideas on


DNA components, and two other lines of evidence. One was the suggestion of Linus Pauling that DNA might have a helical


structure held in place by hydrogen bonds, and the other was X-ray diffraction photos of DNA, showing a helical structure with


distance between the coils, taken by Franklin and Wilkins.


< /p>


直到


40


年代末


50


年代初,


研究者在寻求


DNA


结构过程中,


确立了


Chargaff


的观点和


, Levene


的组成理论 以及其他两


个线索。


一个是


Linus Pauling


的假设,


DNA


可能具 有螺旋结构,


通过氢键连接。


另一个是


X-


衍射图片,


Franklin and Wilkins


提供。



8


/


20


Based on this information Watson and Crick proposed the double helix model of DNA-A twisted ladder-like molecule with two


outer sugar phosphate chains and rungs formed by nucleotide pairs. Paired nucleotides, which always occur as A-T or G-C, are


linked by hydrogen bonds. Watson and Crick also proposed that genetic information is encoded by the sequence of base pairs


along the DNA molecule.



基于这些信息,


Watson




Crick


提出了双螺旋结构模型,成对的核苷酸通过氢键相连,遗传信息就贮藏在碱基对中。



How DNA Replicates



In their model of DNA structure and function, Watson and Crick hypothesized that DNA replicates itself by


the


hydrogen


bonds


joining


A


to


T


and


C


to


G


.


This


process


would


produce


two


opposite


halves


that


could


then


serve


as


templates


for


the


construction


of


new,


complementary


strands. This


model


of


semiconservative


replication


conservative


because each new molecule has one half of the former parent molecule-was later confirmed by the work of



DNA


进行复制是以拉链方式自我复制,产生的两个二分体分别为模板生成互补链,即半保留复制。并由


Meselson




Stahl


验证。



In E. coli DNA replication begins with the formation of a bubblelike structure on the circular chromosome that is produced by


replication forks. Studies of bacterial DNA replication have shown that a growing DNA chain lengthens only in the 5' to 3'


direction (from the 5' carbon of one sugar to the 3' carbon of the next). The leading strand is synthesized continuously, while


the


lagging


strand


is


synthesized


in


short


stretches


known


as


Okazaki


fragments. The


enzyme


DNA


polymerase


links


free


nucleotides as they line up on the template formed by the original strand of the parent molecule.



In eukaryotes DNA replication follows the same general principles as in prokarotes. On the long DNA molecules replication


proceeds (in two directions at once) from hundreds or thousands of points of origin.



大肠杆菌复制开 始时形成泡样复制叉,链生长方向由


5




3



端,前导链连续生成,后随链 由冈崎片段组成,由


DNA



合酶催化 。真核生物复制与原核生物复制相似,但有几百到几千个复制原点(原核一般只有一个复制原点)





第六课



A Home for Life: Formation of the Solar System and Planet Earth



The story of life's origins begins with the formation of the earth. The sequence of events that gave rise to our planet began, in


turn, with the cosmic explosion physicists call the Big Bang. The sun at the center of our solar system condensed from a cloud


of primordial matter roughly 5 billion years ago; the planets, including the earth, condensed about 4.6 billion years ago. The


earth is composed of a number of layers: a solid crust, a semisolid mantle, and a largely molten (liquid) core that has a solid


center. Basic


physical


features


of


Earth


that


may


have


made


the


emergence


of


life


possible


include


the


planet's


size,


temperature, composition, and distance from the sun. The major current hypothesis holds that life arose spontaneously on the


early earth by means of chemical evolution from nonliving substances.



生命起源于地球的形成。大爆炸是我 们行星形成的开始。太阳在


50


亿年前生成,位于太阳系中心, 行星,包括地球,



46


亿年前生成。 地球由多层组成:坚硬的地壳,半流体的地幔,一个很大的溶解中心中存在一个坚实的中心。地球


的基本特征使生命起源成为可能,包括行星的大小,温度,组成以及离太阳的距离。当前主要假设认为, 由非生命的


化学物质进化过程自发产生了生命。



The Emergence of life: Organic and Biological Molecules on a Primitive Planet



Evidence


for


prelife


stages


of


chemical


organization


comes


from


laboratory


experiments


that


try


to


duplicate


the


physical


environment and chemical resources of the early earth. These experiments, including the pioneering work of Miller and Urey,


have successfully produced organic monomers including amino acids, simple sugars, and nucleic acid bases. The probable next


step toward life was the spontaneous linking of such monomers into polymers such as proteinoids and nuclei acids. Current


research suggests that likely sites for this polymerization were clay or rock surfaces.



通过创造出地球早期的自然条件和化学资 源条件,科学家在实验室中已经获得了化学有机体生命前阶段的证据。这些


实验包括米勒 等早期所做工作,成功地产出了有机单体,包括氨基酸,单糖,核酸碱基。这些单体自发的连接成多聚

< p>
体,如类蛋白和核酸,使进入生命状态成为可能。这些聚合作用可能发生在泥土或岩石表面。



Researchers


have


found


that,


when


energy


is


available


to


a


system,


they


can


generate


three


kinds


of


organic


molecular


aggregates. The


Russian


Aleksandr


Oparin


obtained


polymer-rich


droplets,


called


coacervates


from


solutions


of


polymers. Sidney Fox generated proteinoid microspheres from mixtures of amino acids and water. A third laboratory structure


is the liposome, a spherical lipid bilayer that forms from phospholipids. A structure similar to one or more of these aggregates


may have been the precursor of true cells.



9


/


20


当一个系统获得能量时,可 能发生


3


种有机分子的聚集。从多聚物的溶液中俄国科学家获得 了富含小液滴的多聚物,


即凝聚物。从氨基酸和水的混合液中


F ox


获得了类蛋白微球体。实验室第三个结构物质是脂质体,即由磷脂形成的球


形脂双层结构。



Further


steps


in


the


appearance


of


cells


on


the earth


included


the


development


of


RNA


and DNA


as biological


information


molecules. Evidence suggests that RNA, which can form spontaneously under conditions mimicking those of the early earth,


was the first informational molecule. The discovery of RNA ribozymes-RNA that can act as an enzymelike catalyst suggests


that such catalytic RNA also could have assembled new RNAs from early nucleotides. Certain catalytic RNAs can also carry


out sexlike exchanges of pieces of RNA.



RNA


可能是地球早 期形成的第一个贮藏信息的物质。在实验室模拟早期地球自然条件下其可自发形成。核酶的发现说


明它可以催化早期核酸形成新的


RNA


。对于


RNA


物种而言,某些催化


RNA


具备类似于性交换的功能。



Following the development of a lipid-protein surface layer and replicating RNA and DNA informational molecules, the events


leading to the emergence of living cells would have included the origin of the genetic code; the sequestering of RNA or DNA


into cell-like structures; and the development of metabolic pathways.



接下来脂蛋白表层的形成,


RNA


的复制


,DNA


信息 分子的形成,最终导致活细胞的出现,包括最初的遗传密码,


RNA


DNA


被包裹进细胞样的结构中;及代谢途径的建立。< /p>



The Earliest Cells



The oldest fossils that may represent living cells are found in rocks that are about 3.5 billion years old. The cells were probably


anaerobic heterotrophs, with autotrophs arising much later. The first autotrophs produced their own nutrients and released O,-a


metabolic by-product that had a crucial impact on later life forms. The resulting ozone layer in the earth's atmosphere reduced


the penetration of ultraviolet light. As a result, cells would survive in shallow water and on the land surface. The increasing


quantity of atmospheric oxygen also permitted the evolution of aerobic cells and cellular respiration, which in turn signaled the


beginning of the global carbon gh the earliest cells were all prokaryotes, by about 1.5 billion years ago eukaryotes


appeared.



能说明活细胞存 在的最古老化石大约有


35


亿年了。最早出现的细胞可能是厌氧 异氧生物,自养生物很久后出现。最早


的自养生物自己生产营养并释放氧气,这个新陈代 谢副产品对后期生命的形成有一个深远的影响。臭氧层的出现减少


了紫外线的渗透。结果 ,细胞就能够在浅水区和陆地上生存了。大气中氧气数量的增加使得需氧细胞进化并产生细胞

呼吸,这预示着全球碳循环的开始。最早期的细胞都是原核生物,直到


15


亿年后,真核细胞才出现。



The Changing Face of planet Earth



Changes in land masses, the seas, and climate have greatly affected the evolution of life on the earth. The basic parts of the


planet include a light, solid crust over a hot, semisolid mantle and an inner, partially molten core. Massive segments or plates


of


the


crust


move


over


the


mantle


in


the


process


of


continental


drift. Over


the


past


500


million


years,


continental


drift


has


sculpted the earth's crusts to produce the form and distribution of present-day continents. Climatic changes that greatly affected


living organisms accompanied these plate movements; the period was marked by occasional waves of mass extinctions of living


creatures. Organisms were also affected by periods of glaciation that followed variations in the earth's orbit and in the output of


energy by the sun.



大陆板快,海洋的改变,以及气候对地球生命的进化都有深远影响。地幔上大板快地壳的挤压形成大陆 漂移。大陆漂


移雕塑了地壳的外观,使现在大陆形成。伴随板块运动,气候改变对活有机 体有深远影响。在特定时期的生物大量灭


亡高峰是这个时期的见证。生物也受冰河期影响 ,在冰河期,地球轨道和太阳能的输出都发生了很大变化。



Taxonomy: Categorizing the Variety of living Things



Biologists use the binomial system of nomenclature developed by Linnaeus to categorize the varieties of life on the earth. The


system


assigns


each


type


of


organism


to


a


genus


and


species. Organisms


are


then


further


classified


into


higher


taxonomic


categories-family, order, class, division (plants), phylum (animals), and kingdom. Evidence from many subfields of biology,


such


as


biochemistry


and


comparative


anatomy,


helps


define


species


and


higher


taxa


(taxon). And


whereas


species


were


originally


defined


in


terms


of


morphological


traits,


today


biologists


generally


use


the


criterion


of


a


reproductively


isolated


population.



生物学家利 用林奈发展的双名法对生物分类。系统选定每个类型的生物进入属和种,然后将生物进一步分类更高级类


别中,即科,目,纲,门,界。来自于生物化学和比较解剖学等亚生物学领域的证据有助于划分物 种和更高级的分类


单位,然而物种最初依据形态学特征进行分类的,今天生物学家大体上 使用孤立多产的群体作为标准。



Taxonomy


reveals


a


great


deal


about


the


evolutionary


relationships


among


organisms.


A


clade


is


a


taxonomic


unit


whose


10


/


20

-


-


-


-


-


-


-


-



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