Home      Discussion      Topics      Dictionary      Almanac
Signup       Login
Endomembrane system

Endomembrane system

Overview
The endomembrane system is composed of the different membranes that are suspended in the cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...

 within a eukaryotic cell
Eukaryotic Cell
Eukaryotic Cell is an academic journal published by the American Society for Microbiology. The title is commonly abbreviated EC and the ISSN is 1535-9778 for the print version, and 1535-9786 for the electronic version....

. These membranes divide the cell into functional and structural compartments, or organelles. In eukaryote
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

s the organelles of the endomembrane system include: the nuclear envelope
Nuclear envelope
A nuclear envelope is a double lipid bilayer that encloses the genetic material in eukaryotic cells. The nuclear envelope also serves as the physical barrier, separating the contents of the nucleus from the cytosol...

, the endoplasmic reticulum
Endoplasmic reticulum
The endoplasmic reticulum is an organelle of cells in eukaryotic organisms that forms an interconnected network of tubules, vesicles, and cisternae...

, the Golgi apparatus
Golgi apparatus
The Golgi apparatus is an organelle found in most eukaryotic cells. It was identified in 1898 by the Italian physician Camillo Golgi, after whom the Golgi apparatus is named....

, lysosome
Lysosome
thumb|350px|Schematic of typical animal cell, showing subcellular components. [[Organelle]]s: [[nucleoli]] [[cell nucleus|nucleus]] [[ribosomes]] [[vesicle |vesicle]] rough [[endoplasmic reticulum]]...

s, vacuole
Vacuole
A vacuole is a membrane-bound organelle which is present in all plant and fungal cells and some protist, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain...

s, vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

, and the cell membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

. The system is defined more accurately as the set of membranes that form a single functional and developmental unit, either being connected together directly, or exchanging material through vesicle transport.
Discussion
Ask a question about 'Endomembrane system'
Start a new discussion about 'Endomembrane system'
Answer questions from other users
Full Discussion Forum
 
Unanswered Questions
Recent Discussions
Encyclopedia
The endomembrane system is composed of the different membranes that are suspended in the cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...

 within a eukaryotic cell
Eukaryotic Cell
Eukaryotic Cell is an academic journal published by the American Society for Microbiology. The title is commonly abbreviated EC and the ISSN is 1535-9778 for the print version, and 1535-9786 for the electronic version....

. These membranes divide the cell into functional and structural compartments, or organelles. In eukaryote
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

s the organelles of the endomembrane system include: the nuclear envelope
Nuclear envelope
A nuclear envelope is a double lipid bilayer that encloses the genetic material in eukaryotic cells. The nuclear envelope also serves as the physical barrier, separating the contents of the nucleus from the cytosol...

, the endoplasmic reticulum
Endoplasmic reticulum
The endoplasmic reticulum is an organelle of cells in eukaryotic organisms that forms an interconnected network of tubules, vesicles, and cisternae...

, the Golgi apparatus
Golgi apparatus
The Golgi apparatus is an organelle found in most eukaryotic cells. It was identified in 1898 by the Italian physician Camillo Golgi, after whom the Golgi apparatus is named....

, lysosome
Lysosome
thumb|350px|Schematic of typical animal cell, showing subcellular components. [[Organelle]]s: [[nucleoli]] [[cell nucleus|nucleus]] [[ribosomes]] [[vesicle |vesicle]] rough [[endoplasmic reticulum]]...

s, vacuole
Vacuole
A vacuole is a membrane-bound organelle which is present in all plant and fungal cells and some protist, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain...

s, vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

, and the cell membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

. The system is defined more accurately as the set of membranes that form a single functional and developmental unit, either being connected together directly, or exchanging material through vesicle transport. Importantly, the endomembrane system does not include the membranes of mitochondria, chloroplasts or peroxisomes.

The nuclear envelope is a membrane containing two layers, that encompasses the contents of the nucleus. The endoplasmic reticulum (ER) is a synthesis and transport organelle that branches into the cytoplasm in plant and animal cells. The Golgi apparatus is a series of multiple compartments where molecules are packaged for delivery to other cell components or for secretion from the cell. Vacuoles, which are found in both plant and animal cells (though much bigger in plant cells), are responsible for maintaining the shape and structure of the cell as well as storing waste products. A vesicle is a relatively small, membrane-enclosed sac that stores or transports substances. The plasma membrane, also referred to as the cell membrane, is a protective barrier that regulates what enters and leaves the cell. There is also an organelle known as the spitzenkörper
Spitzenkörper
The Spitzenkörper is a structure found in fungal hyphae which is the organizing center for hyphal growth and morphogenesis. It consists of many small vesicles and is present in growing hyphal tips, during spore germination and where branch formation occurs. Its position in the hyphal tip correlates...

 that is only found in fungi, and is connected with hyphal tip growth.

In prokaryote
Prokaryote
The prokaryotes are a group of organisms that lack a cell nucleus , or any other membrane-bound organelles. The organisms that have a cell nucleus are called eukaryotes. Most prokaryotes are unicellular, but a few such as myxobacteria have multicellular stages in their life cycles...

s endomembranes are rare, although in many photosynthetic bacteria the plasma membrane is highly folded and most of the cell cytoplasm is filled with layers of light-gathering membrane. These light-gathering membranes may even form enclosed structures called chlorosome
Chlorosome
A Chlorosome is a photosynthetic antenna complex found in green sulfur bacteria and some green filamentous anoxygenic phototrophs . They differ from other antenna complexes by their large size and lack of protein matrix supporting the photosynthetic pigments...

s in green sulfur bacteria
Green sulfur bacteria
The green sulfur bacteria are a family of obligately anaerobic photoautotrophic bacteria. Most closely related to the distant Bacteroidetes, they are accordingly assigned their own phylum....

.

The organelles of the endomembrane system are related through direct contact or by the transfer of membrane segments as vesicles. Despite these relationships, the various membranes are not identical in structure and function. The thickness, molecular composition, and metabolic behavior of a membrane are not fixed, they may be modified several times during the membrane's life. One unifying characteristic the membranes share is a lipid bilayer
Lipid bilayer
The lipid bilayer is a thin membrane made of two layers of lipid molecules. These membranes are flat sheets that form a continuous barrier around cells. The cell membrane of almost all living organisms and many viruses are made of a lipid bilayer, as are the membranes surrounding the cell nucleus...

, with proteins attached to either side or traversing them.

History of the concept


Most lipids are synthesized in yeast either in the endoplasmic reticulum, lipid particles, or the mitochondrion, with little or no lipid synthesis occurring in the plasma membrane or nuclear membrane. Sphingolipid
Sphingolipid
Sphingolipids are a class of lipids containing a backbone of sphingoid bases, a set of aliphatic amino alcohols that includes sphingosine. They were discovered in brain extracts in the 1870s and were named for the mythological Sphinx because of their enigmatic nature. These compounds play...

 biosynthesis begins in the endoplasmic reticulum, but is completed in the Golgi apparatus. The situation is similar on mammals, with the exception of the first few steps in ether lipid
Ether lipid
Ether lipids are lipids in which one or more of the carbon atoms on glycerol is bonded to an alkyl chain via an ether linkage, as opposed to the usual ester linkage.-Types:...

 biosynthesis, which occur in peroxisomes. The various membranes that enclose the other subcellular organelles must therefore be constructed by transfer of lipids from these sites of synthesis. However, although it is clear that lipid transport is a central process in organelle biogenesis, the mechanisms by which lipids are transported through cells remain poorly understood.

The first proposal that the membranes within cells form a single system that exchanges material between its components was by Morré and Mollenhauer in 1974. This proposal was made as a way of explaining how the various lipid membranes are assembled in the cell, with these membranes being assembled through lipid flow from the sites of lipid synthesis. The idea of lipid flow through a continuous system of membranes and vesicles was an alternative to the various membranes being independent entities that are formed from transport of free lipid components, such as fatty acid
Fatty acid
In chemistry, especially biochemistry, a fatty acid is a carboxylic acid with a long unbranched aliphatic tail , which is either saturated or unsaturated. Most naturally occurring fatty acids have a chain of an even number of carbon atoms, from 4 to 28. Fatty acids are usually derived from...

s and sterol
Sterol
Sterols, also known as steroid alcohols, are a subgroup of the steroids and an important class of organic molecules. They occur naturally in plants, animals, and fungi, with the most familiar type of animal sterol being cholesterol...

s, through the cytosol. Importantly, the transport of lipids through the cytosol and lipid flow through a continuous endomembrane system are not mutually exclusive processes and both may occur in cells.

Nuclear envelope



The nuclear envelope
Nuclear envelope
A nuclear envelope is a double lipid bilayer that encloses the genetic material in eukaryotic cells. The nuclear envelope also serves as the physical barrier, separating the contents of the nucleus from the cytosol...

 surrounds the nucleus
Cell nucleus
In cell biology, the nucleus is a membrane-enclosed organelle found in eukaryotic cells. It contains most of the cell's genetic material, organized as multiple long linear DNA molecules in complex with a large variety of proteins, such as histones, to form chromosomes. The genes within these...

, separating its contents from the cytoplasm. It has two membranes, each a lipid bilayer
Lipid bilayer
The lipid bilayer is a thin membrane made of two layers of lipid molecules. These membranes are flat sheets that form a continuous barrier around cells. The cell membrane of almost all living organisms and many viruses are made of a lipid bilayer, as are the membranes surrounding the cell nucleus...

 with associated proteins. The outer nuclear membrane is constant with the rough endoplasmic reticulum membrane, and like that structure, features ribosomes attached to the surface. The outer membrane is also constant with the inner nuclear membrane since the two layers are fused together at numerous tiny holes called nuclear pore
Nuclear pore
Nuclear pores are large protein complexes that cross the nuclear envelope, which is the double membrane surrounding the eukaryotic cell nucleus. There are about on average 2000 nuclear pore complexes in the nuclear envelope of a vertebrate cell, but it varies depending on cell type and the stage in...

s that perforate the nuclear envelope. These pores are about 120 nm
Nanometre
A nanometre is a unit of length in the metric system, equal to one billionth of a metre. The name combines the SI prefix nano- with the parent unit name metre .The nanometre is often used to express dimensions on the atomic scale: the diameter...

 in diameter and regulate the passage of molecules between the nucleus and cytoplasm, permitting some to pass through the membrane, but not others. Since the nuclear pores are located in an area of high traffic, they play an important role in the physiology
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

 of cells. The space between the outer and inner membranes is called the perinuclear space and is joined with the lumen of the rough ER.

The nuclear envelope's structure is determined by a network of intermediate filaments (protein filaments). This network is organized into lining similar to mesh called the nuclear lamina
Nuclear lamina
The nuclear lamina is a dense fibrillar network inside the nucleus of a eukaryotic cell. It is composed of intermediate filaments and membrane associated proteins. Besides providing mechanical support, the nuclear lamina regulates important cellular events such as DNA replication and cell division...

, which binds to chromatin
Chromatin
Chromatin is the combination of DNA and proteins that make up the contents of the nucleus of a cell. The primary functions of chromatin are; to package DNA into a smaller volume to fit in the cell, to strengthen the DNA to allow mitosis and meiosis and prevent DNA damage, and to control gene...

, integral membrane proteins, and other nuclear components along the inner surface of the nucleus. The nuclear lamina is thought to help materials inside the nucleus reach the nuclear pores and in the disintegration of the nuclear envelope during mitosis
Mitosis
Mitosis is the process by which a eukaryotic cell separates the chromosomes in its cell nucleus into two identical sets, in two separate nuclei. It is generally followed immediately by cytokinesis, which divides the nuclei, cytoplasm, organelles and cell membrane into two cells containing roughly...

 and its reassembly at the end of the process.

The nuclear pores are highly efficient at selectively allowing the passage of materials to and from the nucleus, because the nuclear envelope has a considerable amount of traffic. RNA
RNA
Ribonucleic acid , or RNA, is one of the three major macromolecules that are essential for all known forms of life....

 and ribosomal subunits must be continually transferred from the nucleus to the cytoplasm. Histone
Histone
In biology, histones are highly alkaline proteins found in eukaryotic cell nuclei that package and order the DNA into structural units called nucleosomes. They are the chief protein components of chromatin, acting as spools around which DNA winds, and play a role in gene regulation...

s, gene regulatory proteins, DNA
DNA polymerase
A DNA polymerase is an enzyme that helps catalyze in the polymerization of deoxyribonucleotides into a DNA strand. DNA polymerases are best known for their feedback role in DNA replication, in which the polymerase "reads" an intact DNA strand as a template and uses it to synthesize the new strand....

 and RNA polymerase
RNA polymerase
RNA polymerase is an enzyme that produces RNA. In cells, RNAP is needed for constructing RNA chains from DNA genes as templates, a process called transcription. RNA polymerase enzymes are essential to life and are found in all organisms and many viruses...

s, and other substances essential for nuclear activities must be imported from the cytoplasm. The nuclear envelope of a typical mammalian cell contains 3000–4000 pore complexes. If the cell is synthesizing DNA each pore complex needs to transport about 100 histone molecules per minute. If the cell is growing rapidly, each complex also needs to transport about 6 newly assembled large and small ribosomal subunits per minute from the nucleus to the cytosol, where they are used to synthesize proteins.

Endoplasmic reticulum




The endoplasmic reticulum
Endoplasmic reticulum
The endoplasmic reticulum is an organelle of cells in eukaryotic organisms that forms an interconnected network of tubules, vesicles, and cisternae...

 (ER) is a membranous synthesis and transport organelle that is an extension of the nuclear envelope. More than half the total membrane in eukaryotic cells is accounted for by the ER. The ER is made up of flattened sacs and branching tubules that are thought to interconnect, so that the ER membrane forms a continuous sheet enclosing a single internal space. This highly convoluted space is called the ER lumen and is also referred to as the ER cisternal space. The lumen takes up about ten percent of the entire cell volume. The endoplasmic reticulum membrane allows molecules to be selectively transferred between the lumen and the cytoplasm, and since it is connected to the nuclear envelope, it provides a channel between the nucleus and the cytoplasm.

The ER has a central role in producing, processing, and transporting biochemical compounds
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...

 for use inside and outside of the cell. Its membrane is the site of production of all the transmembrane proteins and lipids for most of the cell's organelles, including the ER itself, the Golgi apparatus, lysosomes, endosomes, mitochondria
Mitochondrion
In cell biology, a mitochondrion is a membrane-enclosed organelle found in most eukaryotic cells. These organelles range from 0.5 to 1.0 micrometers in diameter...

, peroxisome
Peroxisome
Peroxisomes are organelles found in virtually all eukaryotic cells. They are involved in the catabolism of very long chain fatty acids, branched chain fatty acids, D-amino acids, polyamines, and biosynthesis of plasmalogens, etherphospholipids critical for the normal function of mammalian brains...

s, secretory vesicles, and the plasma membrane. Furthermore, almost all of the proteins that will exit the cell, plus those destined for the lumen of the ER, Golgi apparatus, or lysosomes, are originally delivered to the ER lumen. Consequently, many of the proteins found in the cisternal space of the endoplasmic reticulum lumen are there only temporarily as they pass on their way to other locations. Other proteins, however, constantly remain in the lumen and are known as endoplasmic reticulum resident proteins. These special proteins contain a specialized retention signal made up of a specific sequence of amino acids that enables them to be retained by the organelle. An example of an important endoplasmic reticulum resident protein is the chaperon protein known as BiP
Binding immunoglobulin protein
Binding immunoglobulin protein also known as 78 kDa glucose-regulated protein or heat shock 70 kDa protein 5 is a HSP70 molecular chaperone located in the lumen of the endoplasmic reticulum that binds newly-synthesized proteins as they are translocated into the ER, and maintains them in a state...

 which identifies other proteins that have been improperly built or processed and keeps them from being sent to their final destinations.

There are two distinct, though connected, regions of ER that differ in structure and function: smooth ER and rough ER. The rough endoplasmic reticulum is so named because the cytoplasmic surface is covered with ribosomes, giving it a bumpy appearance when viewed through an electron microscope
Electron microscope
An electron microscope is a type of microscope that uses a beam of electrons to illuminate the specimen and produce a magnified image. Electron microscopes have a greater resolving power than a light-powered optical microscope, because electrons have wavelengths about 100,000 times shorter than...

. The smooth ER appears smooth since its cytoplasmic surface lacks ribosomes.

Functions of the smooth ER


In the great majority of cells, smooth ER regions are scarce and are often partly smooth and partly rough. They are sometimes called transitional ER because they contain ER exit sites from which transport vesicles carrying newly synthesized proteins and lipids bud off for transport to the Golgi apparatus. In certain specialized cells, however, the smooth ER is abundant and has additional functions. The smooth ER of theses specialized cells function in diverse metabolic processes, including synthesis of lipids, metabolism of carbohydrates
Carbohydrate metabolism
Carbohydrate metabolism denotes the various biochemical processes responsible for the formation, breakdown and interconversion of carbohydrates in living organisms....

, and detoxification of drugs and poisons.

Enzymes of the smooth ER are vital to the synthesis of lipids, including oils, phospholipids, and steroids. Sex hormones of vertebrates and the steroid hormones secreted by the adrenal glands are among the steroids produced by the smooth ER in animal cells. The cells that synthesis these hormones are rich in smooth ER.

Liver
Liver
The liver is a vital organ present in vertebrates and some other animals. It has a wide range of functions, including detoxification, protein synthesis, and production of biochemicals necessary for digestion...

 cells are another example of specialized cells that contain an abundance of smooth ER. These cells provide an example of the role of smooth ER in carbohydrate metabolism. Liver cells store carbohydrates in the form of glycogen
Glycogen
Glycogen is a molecule that serves as the secondary long-term energy storage in animal and fungal cells, with the primary energy stores being held in adipose tissue...

. The hydrolysis of glycogen leads to the release of glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

 from the liver cells, which is important in the regulation of sugar concentration in the blood. However, glycogen hydrolysis requires glucose phosphate, an ionic form of the sugar that can not exit the cell. An enzyme of the liver cell's smooth ER removes the phosphate from the glucose, so that it can then leave the cell.

Enzymes of the smooth ER can also help detoxify drugs and poisons. Detoxification usually involves the addition of a hydroxyl group to a drug, making the drug more soluble and thus easier to purge from the body. One extensively studied detoxification reaction is carried out by the cytochrome P450 family of enzymes, which catalyze water-insoluble drugs or metabolites that would otherwise accumulate to toxic levels in cell membrane.

Muscle cells have another specialized function of smooth ER. The ER membrane pumps calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

 ions from the cytosol into the cisternal space. When a muscle cell becomes stimulated by a nerve impulse, calcium goes back across the ER membrane into the cytosol and generates the contraction of the muscle cell.

Functions of the rough ER


Many types of cells export proteins produced by ribosomes attached to the rough ER. The ribosomes assemble amino acids into protein units, which are carried into the rough ER for further adjustments. These proteins may be either transmembrane proteins, which become embedded in the membrane of the endoplasmic reticulum, or water-soluble proteins, which are able to pass through the membrane into the lumen. Those that reach the inside of the endoplasmic reticulum are folded into the correct three-dimensional conformation. Chemicals, such as carbohydrates or sugars, are added, then the endoplasmic reticulum either transports the completed proteins, called secretory proteins, to areas of the cell where they are needed, or they are sent to the Golgi apparatus for further processing and modification.

Once secretory proteins are formed, the ER membrane separates them from the proteins that will remain in the cytosol. Secretory proteins depart from the ER enfolded in the membranes of vesicles that bud like bubbles from the transitional ER. These vesicles in transit to another part of the cell are called transport vesicles
Vesicular transport protein
A vesicular transport protein is a transmembrane or membrane associated protein. It regulates or facilitates the movement by vesicles of the contents of the cell.-Types:Examples include:* Archain* ARFs* Clathrin* Caveolin...

. An alternative mechanism for transport of lipids and proteins out of the ER are through lipid transfer proteins at regions called membrane contact site
Membrane contact site
Membrane contact sites are areas within a cell where the membranes of two organelles are arranged in close proximity . These sites are thought to be important in the transport of small molecules such as lipids between organelles, and offer an alternative to the classical mechanism of lipid...

s where the ER becomes closely and stably associated with the membranes of other organelles, such as the plasma membrane, Golgi or lysosomes.

In addition to making secretory proteins, the rough ER makes membranes that grows in place from the addition of proteins and phospholipids. As polypeptides intended to be membrane proteins grow from the ribosomes, they are inserted into the ER membrane itself and are kept there by their hydrophobic portions. The rough ER also produces its own membrane phospholipids; enzymes built into the ER membrane assemble phospholipids. The ER membrane expands and can be transferred by transport vesicles to other components of the endomembrane system.

Golgi apparatus




The Golgi apparatus
Golgi apparatus
The Golgi apparatus is an organelle found in most eukaryotic cells. It was identified in 1898 by the Italian physician Camillo Golgi, after whom the Golgi apparatus is named....

 (also known as the Golgi body and the Golgi complex) is composed of interconnected sacs called cisternae. Its shape can be related to that of a stack of pancakes. The number of these stacks varies with the specific function of the cell. The Golgi apparatus is used by the cell for further protein modification. The section of the Golgi apparatus that receives the vesicles from the ER is known as the cis face, and is usually near the ER. The opposite end of the Golgi apparatus is called the trans face, this is where the modified compounds leave. The trans face is usually facing the plasma membrane, which is where most of the substances the Golgi apparatus modifies are sent.

Vesicles sent off by the ER containing proteins are further altered at the Golgi apparatus and then prepared for secretion from the cell or transport to other parts of the cell. Various things can happen to the proteins on their journey through the enzyme covered space of the Golgi apparatus. The modification and synthesis of the carbohydrate portions of glycoproteins is common in protein processing. The Golgi apparatus removes and substitutes sugar monomers, producing a large variety of oligosaccharides. In addition to modifying proteins, the Golgi also manufactures macromolecules itself. In plant cells, the Golgi produces pectins and other polysaccharides needed by the plant structure.

Once the modification process is completed, the Golgi apparatus sorts the products of its processing and sends them to various parts of the cell. Molecular identification labels or tags are added by the Golgi enzymes to help with this. After everything is organized, the Golgi apparatus sends off its products by budding vesicles from its trans face.

Vacuoles


Vacuole
Vacuole
A vacuole is a membrane-bound organelle which is present in all plant and fungal cells and some protist, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain...

s, like vesicles, are membrane-bounded sacs within the cell. They are larger than vesicles and their specific function varies. The operations of vacuoles are different for plant and animal vacuoles.

In plant cells, vacuoles cover anywhere from 30% to 90% of the total cell volume. Most mature plant cells contain one large central vacuole encompassed by a membrane called the tonoplast. Vacuoles of plant cells act as storage compartments for the nutrients and waste of a cell. The solution that these molecules are stored in is called the cell sap. Pigments that color the cell are sometime located in the cell sap. Vacuoles can also increase the size of the cell, which elongates as water is added, and they control the turgor pressure
Turgor pressure
Turgor Pressure or turgidity is the main pressure of the cell contents against the cell wall in plant cells and bacteria cells, determined by the water content of the vacuole, resulting from osmotic pressure, i.e...

 (the osmotic pressure that keeps the cell wall from caving in). Like lysosomes of animal cells, vacuoles have an acidic pH and contain hydrolytic enzymes. The pH of vacuoles enables them to perform homeostatic procedures in the cell. For example, when the pH in the cells environment drops, the H+ surging into the cytosol can be transferred to a vacuole in order to keep the cytosol's pH constant.

In animals, vacuoles serve in exocytosis
Exocytosis
Exocytosis , also known as 'The peni-cytosis', is the durable process by which a cell directs the contents of secretory vesicles out of the cell membrane...

 and endocytosis
Endocytosis
Endocytosis is a process by which cells absorb molecules by engulfing them. It is used by all cells of the body because most substances important to them are large polar molecules that cannot pass through the hydrophobic plasma or cell membrane...

 processes. Endocytosis refers to when particles are taken into the cell. The material to be taken in is surrounded by the plasma membrane, and then transferred to a vacuole. There are two types of endocytosis, phagocytosis (cell eating) and pinocytosis
Pinocytosis
In cellular biology, pinocytosis is a form of endocytosis in which small particles are brought into the cell—forming an invagination, and then suspended within small vesicles that subsequently fuse with lysosomes to hydrolyze, or to break down, the particles...

 (cell drinking). In phagocytosis, cells engulf large particles such as bacteria. Pinocytosis is the same process, except the substances being ingested are in the fluid form.

Vesicles


Vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

 are small membrane-enclosed transport units that can transfer molecules between different compartments. Most vesicles transfer the membranes assembled in the endoplasmic reticulum to the Golgi apparatus, and then from the Golgi apparatus to various locations.

There are various types of vesicles each with a different protein configuration. Most are formed from specific regions of membranes. When a vesicle buds off from a membrane it contains specific proteins on its cytosolic surface. Each membrane a vesicle travels to contains a marker on its cytosolic surface. This marker corresponds with the proteins on the vesicle traveling to the membrane. Once the vesicle finds the membrane, they fuse.

There are three well known types of vesicles. They are clathrin
Clathrin
Clathrin is a protein that plays a major role in the formation of coated vesicles. Clathrin was first isolated and named by Barbara Pearse in 1975. It forms a triskelion shape composed of three clathrin heavy chains and three light chains. When the triskelia interact they form a polyhedral lattice...

-coated, COPI
COPI
COPI is a protein complex that coats vesicles transporting proteins from the cis end of the Golgi complex back to the rough endoplasmic reticulum , where they were originally synthesized and between golgi compartments. This type of transport is termed as retrograde transport...

-coated, and COPII
COPII
COPII is a type of vesicle coat protein that transports proteins from the rough endoplasmic reticulum to the Golgi apparatus. This is termed anterograde transport. The name "COPII" refers to the specific coat protein complex that initiates the budding process...

-coated vesicles. Each performs different functions in the cell. For example, clathrin-coated vesicles transport substances between the golgi apparatus and the plasma membrane. COPI- and COPII-coated vesicles are frequently used for transportation between the ER and the golgi apparatus.

Lysosomes


Lysosomes are organelles that contain hydrolytic enzymes that are used for intracellular digestion. The main functions of a lysosome are to process molecules taken in by the cell and to recycle worn out cell parts. The enzymes inside of lysosomes are acid hydrolases which require an acidic environment for optimal performance. Lysosomes provide such an environment by maintaining a pH of 5.0 inside of the organelle. If a lysosome were to rupture, the enzymes released would not be very active because of the cytosol's neutral pH. However, if numerous lysosomes leaked the cell could be destroyed from autodigestion.

Lysosomes carry out intracellular digestion by fusing with a vacuole and releasing their enzymes into the vacuole. Through this process, sugars, amino acids, and other monomers pass into the cytosol and become nutrients for the cell. Lysosomes also use their hydrolytic enzymes to recycle the cell's obsolete organelles in a process called autophagy
Autophagy
In cell biology, autophagy, or autophagocytosis, is a catabolic process involving the degradation of a cell's own components through the lysosomal machinery. It is a tightly regulated process that plays a normal part in cell growth, development, and homeostasis, helping to maintain a balance...

. The lysosome engulfs another organelle and uses its enzymes to take apart the ingested material. The resulting organic monomers are then returned to the cytosol for reuse. The last function of a lysosome is to digest the cell itself through autolysis.

Spitzenkörper


The spitzenkörper is a component of the endomembrane system found only in fungi, and is associated with hyphal tip growth
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

. It is a phase-dark body that is composed of an aggregation of membrane-bound vesicles containing cell wall components, serving as a point of assemblage and release of such components intermediate between the Golgi and the cell membrane. The spitzenkörper is motile and generates new hyphal tip growth as it moves forward.


Plasma membrane


The plasma membrane is a phospholipid bilayer membrane that separates the cell from its environment and regulates the transport of molecules and signals into and out of the cell. Embedded in the membrane are proteins that perform the functions of the plasma membrane. The plasma membrane is not a fixed or rigid structure, the molecules that compose the membrane are capable of lateral movement. This movement and the multiple components of the membrane are why it is referred to as a fluid mosaic. Smaller molecules such as carbon dioxide, water, and oxygen can pass through the plasma membrane freely by diffusion
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...

 or osmosis
Osmosis
Osmosis is the movement of solvent molecules through a selectively permeable membrane into a region of higher solute concentration, aiming to equalize the solute concentrations on the two sides...

. Larger molecules needed by the cell are assisted by proteins through active transport
Active transport
Active transport is the movement of a substance against its concentration gradient . In all cells, this is usually concerned with accumulating high concentrations of molecules that the cell needs, such as ions, glucose, and amino acids. If the process uses chemical energy, such as from adenosine...

.

The plasma membrane of a cell has multiple functions. These include transporting nutrients into the cell, allowing waste to leave, preventing materials from entering the cell, averting needed materials from leaving the cell, maintaining the pH of the cytosol, and preserving the osmotic pressure
Osmotic pressure
Osmotic pressure is the pressure which needs to be applied to a solution to prevent the inward flow of water across a semipermeable membrane....

 of the cytosol. Transport proteins which allow some materials to pass through but not others are used for these functions. These proteins use ATP hydrolysis to pump materials against their concentration gradients.

In addition to these universal functions, the plasma membrane has a more specific role in multicellular organisms. Glycoproteins on the membrane assist the cell in recognizing other cells, in order to exchange metabolites and form tissues. Other proteins on the plasma membrane allow attachment to the cytoskeleton
Cytoskeleton
The cytoskeleton is a cellular "scaffolding" or "skeleton" contained within a cell's cytoplasm and is made out of protein. The cytoskeleton is present in all cells; it was once thought to be unique to eukaryotes, but recent research has identified the prokaryotic cytoskeleton...

 and extracellular matrix
Extracellular matrix
In biology, the extracellular matrix is the extracellular part of animal tissue that usually provides structural support to the animal cells in addition to performing various other important functions. The extracellular matrix is the defining feature of connective tissue in animals.Extracellular...

; a function that maintains cell shape and fixes the location of membrane proteins. Enzymes that catalyze reactions are also found on the plasma membrane. Receptor proteins on the membrane have a shape that matches with a chemical messenger, resulting in various cellular responses.

Evolution


The Golgi, ER, and lysosomes are likely to have evolved as a result of the plasma membrane going through invagination
Invagination
Invagination means to fold inward or to sheath. In biology, this can refer to a number of processes.* Invagination is the morphogenetic processes by which an embryo takes form, and is the initial step of gastrulation, the massive reorganization of the embryo from a simple spherical ball of cells,...

. An increase in the overall volume of a cell would require the plasma membrane to fold in order to maintain a constant surface area to volume ratio
Surface area to volume ratio
The surface-area-to-volume ratio also called the surface-to-volume ratio and variously denoted sa/vol or SA:V, is the amount of surface area per unit volume of an object or collection of objects. The surface-area-to-volume ratio is measured in units of inverse distance. A cube with sides of...

. These folds may have led to the specialization of internal membranes to maintain communication with the environment. In the first stages of eukaryotic cell life, the membranes may have been interconnected and attached to the plasma membrane. Later on, as their functions diverged, the membranes may have become separate structures.