Organelle
From Wikipedia, the free encyclopedia
In cell biology, an organelle (pronounced /ɔrɡəˈnɛl/) is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid membrane.
The name organelle comes from the idea that these structures are to cells what an organ is to the body (hence the name organelle, the suffix -elle being a diminutive). Organelles are identified by microscopy, and can also be purified by cell fractionation. There are many types of organelles, particularly in eukaryotic cells. Prokaryotes were once thought not to have organelles, but some examples have now been identified.[1]
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[edit] History and Terminology
In biology, organs are defined as confined functional units within an organism. The analogy of bodily organs to microscopic cellular substructures is obvious, as from even early works, authors of respective textbooks rarely elaborate on the distinction between the two.
Credited as the first[2][3][4] to use a diminutive of organ (i.e. little organ) for cellular structures was German zoologist Karl August Möbius (1884), who used the term "organula" [5] (plural form of organulum, the diminutive of latin organum). From the context, it is clear that he referred to reproduction related structures of protists. In a footnote, which was published as a correction in the next issue of the journal, he justified his suggestion to call organs of unicellular organisms "organella" since they are only differently formed parts of one cell, in contrast to multicellular organs of multicellular organisms. Thus, the original definition was limited to structures of unicellular organisms.
It would take several years before organulum, or the later term organelle, became accepted and expanded in meaning to include subcellular structures in multicellular organisms. Books around 1900 from Valentin Häcker,[6] Edmund Wilson[7] and Oscar Hertwig[8] still referred to cellular organs. Later, both terms came to be used side by side: Bengt Lidforss wrote 1915 (in German) about "Organs or Organells".[9]
Around 1920, the term organelle was used to describe propulsion structures ("motor organelle complex", i.e., flagella and their anchoring)[10] and other protist structures, such as ciliates.[11] Alfred Kühn wrote about centrioles as division organelles, although he stated that, for Vahlkampfias, the alternative 'organelle' or 'product of structural build-up' had not yet been decided, without explaining the difference between the alternatives.[12]
In his 1953 textbook, Max Hartmann used the term for extracellular (pellicula, shells, cell walls) and intracellular skeletons of protists.[13]
Later, the now-widely-used[14][15][16][17] definition of organelle emerged, after which only cellular structures with surrounding membrane had been considered organelles. However, the more original definition of subcellular functional unit in general still coexists.[18][19]
In 1978, Albert Frey-Wyssling suggested that the term organelle should refer only to structures that convert energy, such as centrosomes, ribosomes, and nucleoli.[20][21] This new definition, however, did not win wide recognition.
[edit] Examples
While most cell biologists consider the term organelle to be synonymous with "cell compartment," other cell biologists choose to limit the term organelle to include only those that are DNA-containing, having originated from formerly-autonomous microscopic organisms acquired via endosymbiosis.[22][23][24]
The most notable of these organelles having originated from endosymbiont bacteria are:
- mitochondria (in almost all eukaryotes)
- chloroplasts (in plants, algae and protists).
Other organelles are also suggested to have endosymbiotic origins, (notably the flagellum - see evolution of flagella).
Under the more restricted definition of membrane-bound structures, some parts of the cell do not qualify as organelles. Nevertheless, the use of organelle to refer to non-membrane bound structures such as ribosomes is quite common[25]. This has led some texts to delineate between membrane-bound and non-membrane bound organelles[26]. These structures are large assemblies of macromolecules that carry out particular and specialized functions, but they lack membrane boundaries. Such cell structures include:
[edit] Eukaryotic organelles
Eukaryotes are one of the most structurally complex cell type, and by definition are in part organized by smaller interior compartments, that are themselves enclosed by lipid membranes that resemble the outermost cell membrane. The larger organelles, such as the nucleus and vacuoles, are easily visible with the light microscope. They were among the first biological discoveries made after the invention of the microscope.
Not all eukaryotic cells have every one of the organelles listed below. Exceptional organisms have cells which do not include some organelles that might otherwise be considered universal to eukaryotes (such as mitochondria).[27] There are also occasional exceptions to the number of membranes surrounding organelles, listed in the tables below (e.g., some that are listed as double-membrane are sometimes found with single or triple membranes). In addition, the number of individual organelles of each type found in a given cell varies depending upon the function of that cell.
| Organelle | Main function | Structure | Organisms | Notes |
|---|---|---|---|---|
| chloroplast (plastid) | photosynthesis | double-membrane compartment | plants, protists (rare kleptoplastic organisms) | has some genes; theorized to be engulfed by the ancestral eukaryotic cell (endosymbiosis) |
| endoplasmic reticulum | translation and folding of new proteins (rough endoplasmic reticulum), expression of lipids (smooth endoplasmic reticulum) | single-membrane compartment | all eukaryotes | rough endoplasmic reticulum is covered with ribosomes, has folds that are flat sacs; smooth endoplasmic reticulum has folds that are tubular |
| Golgi apparatus | sorting and modification of proteins | single-membrane compartment | all eukaryotes | cis-face (convex) nearest to rough endoplasmic reticulum; trans-face (concave) farthest from rough endoplasmic reticulum |
| mitochondrion | energy production | double-membrane compartment | most eukaryotes | has some DNA; theorized to be engulfed by the ancestral eukaryotic cell (endosymbiosis) |
| vacuole | storage, homeostasis | single-membrane compartment | eukaryotes | |
| nucleus | DNA maintenance, RNA transcription | double-membrane compartment | all eukaryotes | has bulk of genome |
Mitochondria and chloroplasts, which have double-membranes and their own DNA, are believed to have originated from incompletely consumed or invading prokaryotic organisms, which were adopted as a part of the invaded cell. This idea is supported in the Endosymbiotic theory.
| Organelle/Macromolecule | Main function | Structure | Organisms |
|---|---|---|---|
| acrosome | helps spermatoza fuse with ovum | single-membrane compartment | many animals |
| autophagosome | vesicle which sequesters cytoplasmic material and organelles for degradation | double-membrane compartment | all eukaryotic cells |
| centriole | anchor for cytoskeleton | Microtubule protein | animals |
| cilium | movement in or of external medium; "critical developmental signaling pathway"[28]. | Microtubule protein | animals, protists, few plants |
| eyespot apparatus | detects light, allowing phototaxis to take place | green algae and other unicellular photosynthetic organisms such as euglenids | |
| glycosome | carries out glycolysis | single-membrane compartment | Some protozoa, such as Trypanosomes. |
| glyoxysome | conversion of fat into sugars | single-membrane compartment | plants |
| hydrogenosome | energy & hydrogen production | double-membrane compartment | a few unicellular eukaryotes |
| lysosome | breakdown of large molecules (e.g., proteins + polysaccharides) | single-membrane compartment | most eukaryotes |
| melanosome | pigment storage | single-membrane compartment | animals |
| mitosome | not characterized | double-membrane compartment | a few unicellular eukaryotes |
| myofibril | muscular contraction | bundled filaments | animals |
| nucleolus | ribosome production | protein-DNA-RNA | most eukaryotes |
| parenthesome | not characterized | not characterized | fungi |
| peroxisome | breakdown of metabolic hydrogen peroxide | single-membrane compartment | all eukaryotes |
| ribosome | translation of RNA into proteins | RNA-protein | eukaryotes, prokaryotes |
| vesicle | material transport | single-membrane compartment | all eukaryotes |
Other related structures:
[edit] Prokaryotic organelles
Prokaryotes are not as structurally complex as eukaryotes, and were once thought not to have any internal structures enclosed by lipid membranes. In the past, they were often viewed as having little internal organization; but, slowly, details are emerging about prokaryotic internal structures. An early false turn was the idea developed in the 1970s that bacteria might contain membrane folds termed mesosomes, but these were later shown to be artifacts produced by the chemicals used to prepare the cells for electron microscopy.[30]
However, more recent research has revealed that at least some prokaryotes have microcompartments such as carboxysomes. These subcellular compartments are 100 - 200 nm in diameter and are enclosed by a shell of proteins.[1] Even more striking is the description of membrane-bound magnetosomes in bacteria,[31][32] as well as the nucleus-like structures of the Planctomycetes that are surrounded by lipid membranes.[33]
| Organelle/Macromolecule | Main function | Structure | Organisms |
|---|---|---|---|
| carboxysome | carbon fixation | protein-shell compartment | some bacteria |
| chlorosome | photosynthesis | light harvesting complex | green sulfur bacteria |
| flagellum | movement in external medium | protein filament | some prokaryotes and eukaryotes |
| magnetosome | magnetic orientation | inorganic crystal, lipid membrane | magnetotactic bacteria |
| nucleoid | DNA maintenance, transcription to RNA | DNA-protein | prokaryotes |
| plasmid | DNA exchange | circular DNA | some bacteria |
| ribosome | translation of RNA into proteins | RNA-protein | eukaryotes, prokaryotes |
| thylakoid | photosynthesis | photosystem proteins and pigments | mostly cyanobacteria |
Cell (biology)
From Wikipedia, the free encyclopedia
The cell is the structural and functional unit of all known living organisms. It is the smallest unit of an organism that is classified as living, and is often called the building block of life.[1] Some organisms, such as most bacteria, are unicellular (consist of a single cell). Other organisms, such as humans, are multicellular. (Humans have an estimated 100 trillion or 1014 cells; a typical cell size is 10 µm; a typical cell mass is 1 nanogram.) The largest known cell is an unfertilized ostrich egg cell.[2]
In 1835 before the final cell theory was developed, Jan Evangelista Purkyně observed small "granules" while looking at the plant tissue through a microscope. The cell theory, first developed in 1839 by Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more cells, that all cells come from preexisting cells, that vital functions of an organism occur within cells, and that all cells contain the hereditary information necessary for regulating cell functions and for transmitting information to the next generation of cells.[3]
The word cell comes from the Latin cellula, meaning, a small room. The descriptive name for the smallest living biological structure was chosen by Robert Hooke in a book he published in 1665 when he compared the cork cells he saw through his microscope to the small rooms monks lived in.[4]
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[edit] General principles
Each cell is at least somewhat self-contained and self-maintaining: it can take in nutrients, convert these nutrients into energy, carry out specialized functions, and reproduce as necessary. Each cell stores its own set of instructions for carrying out each of these activities.
All cells have several different abilities:[5]
- Reproduction by cell division: (binary fission/mitosis or meiosis).
- Use of enzymes and other proteins coded for by DNA genes and made via messenger RNA intermediates and ribosomes.
- Metabolism, including taking in raw materials, building cell components, converting energy, molecules and releasing by-products. The functioning of a cell depends upon its ability to extract and use chemical energy stored in organic molecules. This energy is released and then used in metabolic pathways.
- Response to external and internal stimuli such as changes in temperature, pH or levels of nutrients.
- Cell contents are contained within a cell surface membrane that is made from a lipid bilayer with proteins embedded in it.
Some prokaryotic cells contain important internal membrane-bound compartments,[6] but eukaryotic cells have a specialized set of internal membrane compartments.
[edit] Anatomy of cells
There are two types of cells: eukaryotic and prokaryotic. Prokaryotic cells are usually independent, while eukaryotic cells are often found in multicellular organisms.
[edit] Prokaryotic cells
The prokaryote cell is simpler than a eukaryote cell, lacking a nucleus and most of the other organelles of eukaryotes. There are two kinds of prokaryotes: bacteria and archaea; these share a similar overall structure.
A prokaryotic cell has three architectural regions:
- on the outside, flagella and pili project from the cell's surface. These are structures (not present in all prokaryotes) made of proteins that facilitate movement and communication between cells;
- enclosing the cell is the cell envelope – generally consisting of a cell wall covering a plasma membrane though some bacteria also have a further covering layer called a capsule. The envelope gives rigidity to the cell and separates the interior of the cell from its environment, serving as a protective filter. Though most prokaryotes have a cell wall, there are exceptions such as Mycoplasma (bacteria) and Thermoplasma (archaea)). The cell wall consists of peptidoglycan in bacteria, and acts as an additional barrier against exterior forces. It also prevents the cell from expanding and finally bursting (cytolysis) from osmotic pressure against a hypotonic environment. Some eukaryote cells (plant cells and fungi cells) also have a cell wall;
- inside the cell is the cytoplasmic region that contains the cell genome (DNA) and ribosomes and various sorts of inclusions. A prokaryotic chromosome is usually a circular molecule (an exception is that of the bacterium Borrelia burgdorferi, which causes Lyme disease). Though not forming a nucleus, the DNA is condensed in a nucleoid. Prokaryotes can carry extrachromosomal DNA elements called plasmids, which are usually circular. Plasmids enable additional functions, such as antibiotic resistance.
[edit] Eukaryotic cells
Organelles:
(1) nucleolus
(2) nucleus
(3) ribosome
(4) vesicle
(5) rough endoplasmic reticulum (ER)
(6) Golgi apparatus
(7) Cytoskeleton
(8) smooth endoplasmic reticulum
(9) mitochondria
(10) vacuole
(11) cytoplasm
(12) lysosome
(13) centrioles within centrosome
Eukaryotic cells are about 10 times the size of a typical prokaryote and can be as much as 1000 times greater in volume. The major difference between prokaryotes and eukaryotes is that eukaryotic cells contain membrane-bound compartments in which specific metabolic activities take place. Most important among these is the presence of a cell nucleus, a membrane-delineated compartment that houses the eukaryotic cell's DNA. It is this nucleus that gives the eukaryote its name, which means "true nucleus." Other differences include:
- The plasma membrane resembles that of prokaryotes in function, with minor differences in the setup. Cell walls may or may not be present.
- The eukaryotic DNA is organized in one or more linear molecules, called chromosomes, which are associated with histone proteins. All chromosomal DNA is stored in the cell nucleus, separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria also contain some DNA.
- Eukaryotes can move using cilia or flagella. The flagella are more complex than those of prokaryotes.
| Prokaryotes | Eukaryotes | |
|---|---|---|
| Typical organisms | bacteria, archaea | protists, fungi, plants, animals |
| Typical size | ~ 1–10 µm | ~ 10–100 µm (sperm cells, apart from the tail, are smaller) |
| Type of nucleus | nucleoid region; no real nucleus | real nucleus with double membrane |
| DNA | circular (usually) | linear molecules (chromosomes) with histone proteins |
| RNA-/protein-synthesis | coupled in cytoplasm | RNA-synthesis inside the nucleus protein synthesis in cytoplasm |
| Ribosomes | 50S+30S | 60S+40S |
| Cytoplasmatic structure | very few structures | highly structured by endomembranes and a cytoskeleton |
| Cell movement | flagella made of flagellin | flagella and cilia containing microtubules; lamellipodia and filopodia containing actin |
| Mitochondria | none | one to several thousand (though some lack mitochondria) |
| Chloroplasts | none | in algae and plants |
| Organization | usually single cells | single cells, colonies, higher multicellular organisms with specialized cells |
| Cell division | Binary fission (simple division) | Mitosis (fission or budding) Meiosis |
| Typical animal cell | Typical plant cell | |
|---|---|---|
| Organelles |
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[edit] Subcellular components
All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, separates its interior from its environment, regulates what moves in and out (selectively permeable), and maintains the electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. All cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules in cells. This article will list these primary components of the cell, then briefly describe their function.
[edit] Cell membrane: A cell's defining boundary
The cytoplasm of a cell is surrounded by a cell membrane or plasma membrane. The plasma membrane in plants and prokaryotes is usually covered by a cell wall. This membrane serves to separate and protect a cell from its surrounding environment and is made mostly from a double layer of lipids (hydrophobic fat-like molecules) and hydrophilic phosphorus molecules. Hence, the layer is called a phospholipid bilayer. It may also be called a fluid mosaic membrane. Embedded within this membrane is a variety of protein molecules that act as channels and pumps that move different molecules into and out of the cell. The membrane is said to be 'semi-permeable', in that it can either let a substance (molecule or ion) pass through freely, pass through to a limited extent or not pass through at all. Cell surface membranes also contain receptor proteins that allow cells to detect external signaling molecules such as hormones.
[edit] Cytoskeleton: A cell's scaffold
The cytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps during endocytosis, the uptake of external materials by a cell, and cytokinesis, the separation of daughter cells after cell division; and moves parts of the cell in processes of growth and mobility. The eukaryotic cytoskeleton is composed of microfilaments, intermediate filaments and microtubules. There is a great number of proteins associated with them, each controlling a cell's structure by directing, bundling, and aligning filaments. The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell shape, polarity and cytokinesis.[7]
[edit] Genetic material
Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). Most organisms use DNA for their long-term information storage, but some viruses (e.g., retroviruses) have RNA as their genetic material. The biological information contained in an organism is encoded in its DNA or RNA sequence. RNA is also used for information transport (e.g., mRNA) and enzymatic functions (e.g., ribosomal RNA) in organisms that use DNA for the genetic code itself. Transfer RNA (tRNA) molecules are used to add specific amino acids during the process of protein translation.
Prokaryotic genetic material is organized in a simple circular DNA molecule (the bacterial chromosome) in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different, linear molecules called chromosomes inside a discrete nucleus, usually with additional genetic material in some organelles like mitochondria and chloroplasts (see endosymbiotic theory).
A human cell has genetic material in the nucleus (the nuclear genome) and in the mitochondria (the mitochondrial genome). In humans the nuclear genome is divided into 23 pairs of linear DNA molecules called chromosomes. The mitochondrial genome is a circular DNA molecule distinct from the nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, it codes for 13 proteins involved in mitochondrial energy production as well as specific tRNAs.
Foreign genetic material (most commonly DNA) can also be artificially introduced into the cell by a process called transfection. This can be transient, if the DNA is not inserted into the cell's genome, or stable, if it is. Certain viruses also insert their genetic material into the genome.
[edit] Organelles
The human body contains many different organs, such as the heart, lung, and kidney, with each organ performing a different function. Cells also have a set of "little organs," called organelles, that are adapted and/or specialized for carrying out one or more vital functions.
There are several types of organelles within an animal cell. Some (such as the nucleus and golgi apparatus) are typically solitary, while others (such as mitochondria, peroxisomes and lysosomes) can be numerous (hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.
- Mitochondria and Chloroplasts – the power generators
- Mitochondria are self-replicating organelles that occur in various numbers, shapes, and sizes in the cytoplasm of all eukaryotic cells. Mitochondria play a critical role in generating energy in the eukaryotic cell. Mitochondria generate the cell's energy by the process of oxidative phosphorylation, utilizing oxygen to release energy stored in cellular nutrients (typically pertaining to glucose) to generate ATP. Mitochondria multiply by splitting in two.
- Organelles that are modified chloroplasts are broadly called plastids, and are involved in energy storage through the process of photosynthesis, which utilizes solar energy to generate carbohydrates and oxygen from carbon dioxide and water.
- Mitochondria and chloroplasts each contain their own genome, which is separate and distinct from the nuclear genome of a cell. Both of these organelles contain this DNA in circular plasmids, much like prokaryotic cells, strongly supporting the evolutionary theory of endosymbiosis; since these organelles contain their own genomes and have other similarities to prokaryotes, they are thought to have developed through a symbiotic relationship after being engulfed by a primitive cell.
- Ribosomes
- The ribosome is a large complex of RNA and protein molecules. They each consist of two subunits, and act as an assembly line where mRNA from the nucleus is used to synthesise proteins from amino acids. Ribosomes can be found either floating freely or bound to a membrane (the rough endoplasmatic reticulum in eukaryotes, or the cell membrane in prokaryotes). [8]
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[edit] Structures outside the cell wall
[edit] Capsule
A gelatinous capsule is present in some bacteria outside the cell wall. The capsule may be polysaccharide as in pneumococci, meningococci or polypeptide as bacillus anthracis or hyaluronic acid as in streptococci. Capsules not marked by ordinary stain and can detected by special stain. The capsule is antigenic. The capsule has antiphagocytic function so it determines the virulence of many bacteria. It also plays a role in attachment of the organism to mucous membranes.
[edit] Flagella
Flagella are the organelles of mobility. They arise from cytoplasm and extrude through the cell wall. They are long and thick thread like appendages, protein in nature, formed of flagellin protein (antigenic). They can not be stained by gram stain. They have a special stain. According to their arrangement they may be monotrichate, amphitrichate, lophotrichate, peritrichate.
[edit] Fimbriae (pili)
They are short and thin hair like filaments, formed of protein called pilin (antigenic). Fimbriae are responsible for attachment of bacteria to specific receptors of human cell (adherence). There are special types of pili called (sex pili) involved in the process of conjunction.
[edit] Cell functions
[edit] Cell growth and metabolism
Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct divisions: catabolism, in which the cell breaks down complex molecules to produce energy and reducing power, and anabolism, in which the cell uses energy and reducing power to construct complex molecules and perform other biological functions. Complex sugars consumed by the organism can be broken down into a less chemically-complex sugar molecule called glucose. Once inside the cell, glucose is broken down to make adenosine triphosphate (ATP), a form of energy, via two different pathways.
The first pathway, glycolysis, requires no oxygen and is referred to as anaerobic metabolism. Each reaction is designed to produce some hydrogen ions that can then be used to make energy packets (ATP). In prokaryotes, glycolysis is the only method used for converting energy.
The second pathway, called the Krebs cycle, or citric acid cycle, occurs inside the mitochondria and is capable of generating enough ATP to run all the cell functions.
Within the nucleus of the cell (light blue), genes (DNA, dark blue) are transcribed into RNA. This RNA is then subject to post-transcriptional modification and control, resulting in a mature mRNA (red) that is then transported out of the nucleus and into the cytoplasm (peach), where it undergoes translation into a protein. mRNA is translated by ribosomes (purple) that match the three-base codons of the mRNA to the three-base anti-codons of the appropriate tRNA. Newly-synthesized proteins (black) are often further modified, such as by binding to an effector molecule (orange), to become fully active.
[edit] Creation of new cells
Cell division involves a single cell (called a mother cell) dividing into two daughter cells. This leads to growth in multicellular organisms (the growth of tissue) and to procreation (vegetative reproduction) in unicellular organisms.
Prokaryotic cells divide by binary fission. Eukaryotic cells usually undergo a process of nuclear division, called mitosis, followed by division of the cell, called cytokinesis. A diploid cell may also undergo meiosis to produce haploid cells, usually four. Haploid cells serve as gametes in multicellular organisms, fusing to form new diploid cells.
DNA replication, or the process of duplicating a cell's genome, is required every time a cell divides. Replication, like all cellular activities, requires specialized proteins for carrying out the job.
[edit] Protein synthesis
Cells are capable of synthesizing new proteins, which are essential for the modulation and maintenance of cellular activities. This process involves the formation of new protein molecules from amino acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two major steps: transcription and translation.
Transcription is the process where genetic information in DNA is used to produce a complementary RNA strand. This RNA strand is then processed to give messenger RNA (mRNA), which is free to migrate through the cell. mRNA molecules bind to protein-RNA complexes called ribosomes located in the cytosol, where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence by binding to transfer RNA (tRNA) adapter molecules in binding pockets within the ribosome. The new polypeptide then folds into a functional three-dimensional protein molecule.
[edit] Cell movement or motility
Cells can move during many processes: such as wound healing, the immune response and cancer metastasis. For wound healing to occur, white blood cells and cells that ingest bacteria move to the wound site to kill the microorganisms that cause infection.
At the same time fibroblasts (connective tissue cells) move there to remodel damaged structures. In the case of tumor development, cells from a primary tumor move away and spread to other parts of the body. Cell motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[9] The process is divided into three steps – protrusion of the leading edge of the cell, adhesion of the leading edge and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each of these steps is driven by physical forces generated by unique segments of the cytoskeleton.[10][11]
[edit] Evolution
The origin of cells has to do with the origin of life, which began the history of life on Earth.
[edit] Origin of the first cell
There are three leading hypotheses for the source of small molecules that would make up life in an early Earth. One is that they came from meteorites (see Murchison meteorite). Another is that they were created at deep-sea vents. A third is that they were synthesized by lightning in a reducing atmosphere (see Miller–Urey experiment); although it is not sure Earth had such an atmosphere. There is essentially no experimental data to tell what the first self-replicate forms were. RNA is generally assumed to be the earliest self-replicating molecule, as it is capable of both storing genetic information and catalyze chemical reactions (see RNA world hypothesis). But some other entity with the potential to self-replicate could have preceded RNA, like clay or peptide nucleic acid.[12]
Cells emerged at least 3.0–3.3 billion years ago. The current belief is that these cells were heterotrophs. An important characteristic of cells is the cell membrane, composed of a bilayer of lipids. The early cell membranes were probably more simple and permeable than modern ones, with only a single fatty acid chain per lipid. Lipids are known to spontaneously form bilayered vesicles in water, and could have preceded RNA. But the first cell membranes could also have been produced by catalytic RNA, or even have required structural proteins before they could form.[13]
[edit] Origin of eukaryotic cells
The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. It is almost certain that DNA-bearing organelles like the mitochondria and the chloroplasts are what remains of ancient symbiotic oxygen-breathing proteobacteria and cyanobacteria, respectively, where the rest of the cell seems to be derived from an ancestral archaean prokaryote cell – a theory termed the endosymbiotic theory.
There is still considerable debate about whether organelles like the hydrogenosome predated the origin of mitochondria, or viceversa: see the hydrogen hypothesis for the origin of eukaryotic cells.
Sex, as the stereotyped choreography of meiosis and syngamy that persists in nearly all extant eukaryotes, may have played a role in the transition from prokaryotes to eukaryotes. An 'origin of sex as vaccination' theory suggests that the eukaryote genome accreted from prokaryan parasite genomes in numerous rounds of lateral gene transfer. Sex-as-syngamy (fusion sex) arose when infected hosts began swapping nuclearized genomes containing co-evolved, vertically transmitted symbionts that conveyed protection against horizontal infection by more virulent symbionts.[14]
Endosymbiotic theory
From Wikipedia, the free encyclopedia
The endosymbiotic theory concerns the origins of mitochondria and plastids (e.g. chloroplasts), which are organelles of eukaryotic cells. According to this theory, these organelles originated as separate prokaryotic organisms which were taken inside the cell as endosymbionts. Mitochondria developed from proteobacteria (in particular, Rickettsiales or close relatives) and chloroplasts from cyanobacteria.
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[edit] History
The endosymbiotic theory was first articulated by the Russian botanist Konstantin Mereschkowsky in 1905.[1] Mereschkowsky was familiar with work by botanist Andreas Schimper, who had observed in 1883 that the division of chloroplasts in green plants closely resembled that of free-living cyanobacteria, and who had himself tentatively proposed (in a footnote) that green plants had arisen from a symbiotic union of two organisms.[2] Ivan Wallin extended the idea of an endosymbiotic origin to mitochondria in the 1920s.[3] These theories were initially dismissed or ignored. More detailed electron microscopic comparisons between cyanobacteria and chloroplasts (for example studies by Hans Ris[4]), combined with the discovery that plastids and mitochondria contain their own DNA[5] (which by that stage was recognized to be the hereditary material of organisms) led to a resurrection of the idea in the 1960s.
The endosymbiotic hypothesis was popularized by Lynn Margulis. In her 1981 work Symbiosis in Cell Evolution she argued that eukaryotic cells originated as communities of interacting entities, including endosymbiotic spirochaetes that developed into eukaryotic flagella and cilia. This last idea has not received much acceptance, because flagella lack DNA and do not show ultrastructural similarities to prokaryotes. See also Evolution of flagella.
According to Margulis and Sagan,[6] "Life did not take over the globe by combat, but by networking" (i.e., by cooperation)[7].
The possibility that peroxisomes may have an endosymbiotic origin has also been considered, although they lack DNA. Christian de Duve proposed that they may have been the first endosymbionts, allowing cells to withstand growing amounts of free molecular oxygen in the Earth's atmosphere. However, it now appears that they may be formed de novo, contradicting the idea that they have a symbiotic origin (Gabaldón et al. 2006).
It is also believed that these endosymbionts transferred some of their own DNA to the host cell's nucleus during the evolutionary transition from a symbiotic community to an instituted eukaryotic cell. This hypothesis is thought to be possible because it is known today from scientific observation that transfer of DNA occurs between prokaryotic species, even if they are not closely related. Prokaryotes can take up DNA from their surroundings and have a limited ability to incorporate it into their own genome.
[edit] Evidence
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Evidence that mitochondria and plastids arose from ancient endosymbiosis of bacteria is as follows:
- New mitochondria and plastids are formed only through a process similar to binary fission. In some algae, such as Euglena, the plastids can be destroyed by certain chemicals or prolonged absence of light without otherwise affecting the cell. In such a case, the plastids will not regenerate.
- They are surrounded by two or more membranes, and the innermost of these shows differences in composition from the other membranes of the cell. The composition is like that of a prokaryotic cell membrane.
- Both mitochondria and plastids contain DNA that is different from that of the cell nucleus and that is similar to that of bacteria (in being circular in shape and in its size).
- DNA sequence analysis and phylogenetic estimates suggests that nuclear DNA contains genes that probably came from plastids.
- These organelles' ribosomes are like those found in bacteria (70s).
- Proteins of organelle origin, like those of bacteria, use N-formylmethionine as the initiating amino acid.
- Much of the internal structure and biochemistry of plastids, for instance the presence of thylakoids and particular chlorophylls, is very similar to that of cyanobacteria. Phylogenetic estimates constructed with bacteria, plastids, and eukaryotic genomes also suggest that plastids are most closely related to cyanobacteria.
- Mitochondria have several enzymes and transport systems similar to those of prokaryotes.
- Some proteins encoded in the nucleus are transported to the organelle, and both mitochondria and plastids have small genomes compared to bacteria. This is consistent with an increased dependence on the eukaryotic host after forming an endosymbiosis. Most genes on the organellar genomes have been lost or moved to the nucleus. Most genes needed for mitochondrial and plastid function are located in the nucleus. Many originate from the bacterial endosymbiont.
- Plastids are present in very different groups of protists, some of which are closely related to forms lacking plastids. This suggests that if chloroplasts originated de novo, they did so multiple times, in which case their close similarity to each other is difficult to explain.
- Many of these protists contain "secondary" plastids that have been acquired from other plastid-containing eukaryotes, not from cyanobacteria directly.
- Among the eukaryotes that acquired their plastids directly from bacteria (known as Primoplantae), the glaucophyte algae have chloroplasts that strongly resemble cyanobacteria. In particular, they have a peptidoglycan cell wall between their two membranes.
- Mitochondria and plastids are just about the same size as bacteria.
[edit] Secondary endosymbiosis
Primary endosymbiosis involves the engulfment of a bacterium by another free living organism. Secondary endosymbiosis occurs when the product of primary endosymbiosis is itself engulfed and retained by another free living eukaryote. Secondary endosymbiosis has occurred several times and has given rise to extremely diverse groups of algae and other eukaryotes. Some organisms can take opportunistic advantage of a similar process, where they engulf an alga and use the products of its photosynthesis, but once the prey item dies (or is lost) the host returns to a free living state. Obligate secondary endosymbionts become dependent on their organelles and are unable to survive in their absence (for a review see McFadden 2001[8]).
One possible secondary endosymbiosis in process has been observed by Okamoto & Inouye (2005). The heterotrophic protist Hatena behaves like a predator until it ingests a green alga, which loses its flagella and cytoskeleton, while Hatena, now a host, switches to photosynthetic nutrition, gains the ability to move towards light and loses its feeding apparatus.
The process of secondary endosymbiosis left its evolutionary signature within the unique topography of plastid membranes. Secondary plastids are surrounded by three (in euglenophytes and some dinoflagellates) or four membranes (in haptophytes, heterokonts, cryptophytes, and chlorarachniophytes). The two additional membranes are thought to correspond to the plasma membrane of the engulfed alga and the phagosomal membrane of the host cell. The endosymbiotic acquisition of a eukaryote cell is represented in the cryptophytes; where the remnant nucleus of the red algal symbiont (the nucleomorph) is present between the two inner and two outer plastid membranes.[citation needed]
Despite the diversity of organisms containing plastids, the morphology, biochemistry, genomic organisation, and molecular phylogeny of plastid RNAs and proteins suggest a single origin of all extant plastids – although this theory is still debated. [9][10]
[edit] Problems
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- Neither mitochondria nor plastids can survive in oxygen or outside the cell, having lost many essential genes required for survival. The standard counterargument points to the large timespan that the mitochondria/plastids have co-existed with their hosts. In this view, genes and systems which were no longer necessary were simply deleted, or in many cases, transferred into the host genome instead. (In fact these transfers constitute an important way for the host cell to regulate plastid or mitochondrial activity.)
- The transfer of genes from mitochondria and plastids to the “host genome” or cell nucleus raises a further problem: why were all genes not transferred? In other words, why do any genes at all remain in mitochondria and plastids? This problem is addressed by the CoRR Hypothesis which proposes that genes and respiratory chain proteins are Co-located for Redox Regulation.
- A large cell, especially one equipped for phagocytosis, has vast energetic requirements, which cannot be achieved without the internalisation of energy production (due to the decrease in the surface area to volume ratio as size increases). This implies that, for the cell to gain mitochondria, it could not have been a primitive eukaryote, but instead a prokaryotic cell. This in turn implies that the emergence of the eukaryotes and the formation of mitochondria were achieved simultaneously.
- Genetic analysis of small eukaryotes that lack mitochondria shows that they all still retain genes for mitochondrial proteins. This implies that all these eukaryotes once had mitochondria. This objection can be answered if, as suggested above, the origin of the eukaryotes coincided with the formation of mitochondria.
These last two problems are accounted for in the Hydrogen hypothesis.
Symbiogenesis
From Wikipedia, the free encyclopedia
Symbiogenesis is the merging of two separate organisms to form a single new organism. The idea originated with Konstantin Mereschkowsky in his 1926 book Symbiogenesis and the Origin of Species, which proposed that chloroplasts originate from cyanobacteria captured by a protozoan.[1] Today both chloroplasts and mitochondria are believed to have such an origin; this is the endosymbiotic theory.
In Acquiring Genomes: A Theory of the Origins of Species, biologist Dr. Lynn Margulis argued that symbiogenesis is a primary force in evolution. According to her theory, acquisition and accumulation of random mutations are not sufficient to explain how inherited variations occur; rather, new organelles, bodies, organs, and species arise from symbiogenesis.[2] Whereas the classical interpretation of evolution (the modern evolutionary synthesis) emphasizes competition as the main force behind evolution, Margulis emphasizes cooperation.[3]
Many ecologists agree, but this idea has little support from other evolutionary biologists[citation needed]. They see little evidence that symbiogenesis has had a major impact on eukaryotic life, or that much of its diversification can be attributed to it.[citation needed] Other than the two examples of mitochondria and chloroplasts, there is no clear evidence of other major traits or transitions that can be attributed to symbiogenesis.[citation needed]
A fundamental principle of modern evolutionary theory is that mutations arise one at a time and either spread through the population or not, depending on whether they offer an individual fitness advantage. Nevertheless, this general case may not apply to all examples of evolutionary change. Indeed, genome mapping techniques have revealed that family trees of the major taxa appear to be extensively cross-linked—possibly due to lateral gene transfer.[4]
Viral eukaryogenesis is the hypothesis that the cell nucleus of eukaryotic life forms evolved from a large DNA virus in a form of endosymbiosis within an archaea cell, being a form of symbiogenesis. It was proposed by Philip Bell in 2001, and gained support as large complex DNA viruses capable of protein biosynthesis (such as Mimivirus) have been discovered.
A number of precepts in the theory are possible. For instance, a helical virus with a bilipid envelope bears a distinct resemblance to a highly simplified cellular nucleus (ie: a DNA chromosome encapsulated within a lipid membrane). To consider the concept logically, a large DNA virus would take control of a bacterial or archaeal cell. Instead of replicating and destroying the host cell, it would remain within the cell. With the virus in control of the host cell's molecular machinery it would effectively become a "nucleus" of sorts. Through the processes of mitosis and cytokinesis, the virus would thus hijack the entire cell—an extremely favourable way to ensure its survival.
PABPII, or polyadenine binding protein II, is a protein involved in the assembly of the polyadenine tail added to newly synthesized pre-messenger RNA (mRNA) molecules during the process of gene transcription. It is a regulatory protein that controls the rate at which polyadenine polymerase (PAP) adds adenine nucleotides to the 3' end of the growing tail. In the absence of PABPII, PAP adds adenines slowly. PABPII then binds to the short polyadenine tail and induces an acceleration in the rate of addition by PAP until the tail has grown to about 200 adenines long. The mechanism by which PABPII signals the termination of the polymerization reaction once the tail has reached its required length is not clear.
PABPII is distinct from the related protein PABPI in being localized to the cell nucleus rather than the cytoplasm.
Cytoplasm
From Wikipedia, the free encyclopedia
(1) nucleolus
(2) nucleus
(3) ribosomes (little dots)
(4) vesicle
(5) rough endoplasmic reticulum (ER)
(6) Golgi apparatus
(7) Cytoskeleton
(8) smooth ER
(9) mitochondria
(10) vacuole
(11) cytoplasm
(12) lysosome
(13) centrioles within centrosome
The cytoplasm is the part of a cell that is enclosed within the plasma membrane. In eukaryotic cells, the cytoplasm contains organelles, such as mitochondria, which are filled with liquid that is kept separate from the rest of the cytoplasm by biological membranes. The contents of the cell nucleus are not part of the cytoplasm and are instead called the nucleoplasm. The cytoplasm is the site where most cellular activities occur, such as many metabolic pathways like glycolysis, and processes such as cell division. The inner, granular mass is called the endoplasm and the outer, clear and glassy layer is called the cell cortex or the ectoplasm.
The part of the cytoplasm that is not held within organelles is called the cytosol. The cytosol is a complex mixture of cytoskeleton filaments, dissolved molecules, and water that fills much of the volume of a cell. The cytosol is a gel, with a network of fibers dispersed through water. Due to this network of pores and high concentrations of dissolved macromolecules, such as proteins, an effect called macromolecular crowding occurs and the cytosol does not act as an ideal solution. This crowding effect alters how the components of the cytosol interact with each other.
Contents[show] |
[edit] Constituents
The cytoplasm has three major elements; the cytosol, organelles and inclusions.
[edit] Cytosol
The cytosol is the portion of a cell that is not enclosed within membrane-bound organelles. The cytosol is a translucent fluid in which the other cytoplasmic elements are suspended. Cytosol makes up about 70 % of the cell volume and is composed of water, salts and organic molecules.[1] The cytoplasm also contains the protein filaments that make up the cytoskeleton, as well as soluble proteins and large structures such as ribosomes, proteasomes, and the mysterious vault complexes.[2] The inner, granular and more fluid portion of the cytoplasm is referred to as endoplasm.
[edit] Organelles
Organelles are membrane-bound compartments within the cell that have specific functions. Some major organelles that are suspended in the cytosol are the mitochondria, the endoplasmic reticulum, the Golgi apparatus, lysosomes, and in plant cells chloroplasts.
[edit] Cytoplasmic inclusions
The inclusions are small particles of insoluble substances suspended in the cytosol. A huge range of inclusions exist in different cell types, and range from crystals of calcium oxalate or silicon dioxide in plants,[3][4] to granules of energy-storage materials such as starch,[5] glycogen,[6] or polyhydroxybutyrate.[7] A particularly widespread example are lipid droplets, which are spherical droplets composed of lipids and proteins that are used in both prokaryotes and eukaryotes as a way of storing lipids such as fatty acids and sterols.[8] Lipid droplets make up much of the volume of adipocytes, which are specialized lipid-storage cells, but they are also found in a range of other cell types.
