In cellular biology, inclusions are diverse intracellularShively, J. M. (ed.). (2006). ''Microbiology Monographs Vol. 1: Inclusions in Prokaryotes''. Berlin, Heidelberg: Springer-Verlag
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non-living substances ( ergastic substances) that are not bound by membranes. Inclusions are stored nutrients/ deutoplasmic substances, secretory products, and

pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...

granules. Examples of inclusions are

glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. It is the main storage form of glucose in the human body. Glycogen functions as one of three regularly used forms ...

granules in the

liver The liver is a major metabolic organ (anatomy), organ exclusively found in vertebrates, which performs many essential biological Function (biology), functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of var ...

and

muscle cell A muscle cell, also known as a myocyte, is a mature contractile Cell (biology), cell in the muscle of an animal. In humans and other vertebrates there are three types: skeletal muscle, skeletal, smooth muscle, smooth, and Cardiac muscle, cardiac ...

lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...

droplets in fat cells, pigment granules in certain cells of

skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...

and

hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...

, and

crystal A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macros ...

s of various types.Leslie P. Gartner and James L. Hiatt; Text book of Histology; 3rd edition Cytoplasmic inclusions are an example of a biomolecular condensate arising by liquid-solid, liquid-gel or liquid-liquid

phase separation Phase separation is the creation of two distinct Phase (matter), phases from a single homogeneous mixture. The most common type of phase separation is between two immiscible liquids, such as oil and water. This type of phase separation is kn ...

. These structures were first observed by O. F. Müller in 1786.

Examples

Glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. It is the main storage form of glucose in the human body. Glycogen functions as one of three regularly used forms ...

: Glycogen is the most common form of

glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...

in animals and is especially abundant in cells of muscles, and liver. It appears in electron micrograph as clusters, or a rosette of beta particles that resemble

ribosomes Ribosomes () are macromolecular machines, found within all cells, that perform biological protein synthesis (messenger RNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA molecules to fo ...

, located near the smooth endoplasmic reticulum. Glycogen is an important energy source of the cell; therefore, it will be available on demand. The enzymes responsible for glycogenolysis degrade glycogen into individual molecules of glucose and can be utilized by multiple organs of the body.Fawcett; The cell, 2nd edition

Lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...

s: Lipids, which are stored as

triglycerides A triglyceride (from ''wikt:tri-#Prefix, tri-'' and ''glyceride''; also TG, triacylglycerol, TAG, or triacylglyceride) is an ester derived from glycerol and three fatty acids. Triglycerides are the main constituents of body fat in humans and oth ...

, are the common form of inclusions. They are stored not only in specialized cells ( adipocytes) but also are located as individuals droplets in various cell types, especially hepatocytes. These are fluid at body temperature and appear in living cells as refractile spherical droplets. Lipids yield more than twice as many calories per gram as do carbohydrates. On demand, they serve as a local store of energy and a potential source of short carbon chains that are used by the cell in its synthesis of membranes and other lipid containing structural components or secretory products.

Crystal A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macros ...

s: Crystalline inclusions have long been recognized as normal constituents of certain cell types such as Sertoli cells and Leydig cells of the human testis, and are found occasionally in

macrophages Macrophages (; abbreviated MPhi, φ, MΦ or MP) are a type of white blood cell of the innate immune system that engulf and digest pathogens, such as cancer cells, microbes, cellular debris and foreign substances, which do not have proteins that ...

. It is believed that these structures are crystalline forms of certain proteins, and are located everywhere in the cell, including the nucleus,

mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...

endoplasmic reticulum The endoplasmic reticulum (ER) is a part of a transportation system of the eukaryote, eukaryotic cell, and has many other important functions such as protein folding. The word endoplasmic means "within the cytoplasm", and reticulum is Latin for ...

, Golgi body, and free in cytoplasmic matrix.

Pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...

s: The most common pigment in the body, besides

hemoglobin Hemoglobin (haemoglobin, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobin ...

of red blood cells, is

melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...

, manufactured by

melanocytes Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and hea ...

of the skin and hair, pigment cells of the retina and specialized nerve cells in the

substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra a ...

of the brain. These pigments have protective functions in the skin and aid in the sense of sight in the retina, but their function in neurons is not understood completely. Furthermore, cardiac tissue and central nervous system neurons show a yellow to brown pigment called lipofuscin, believed by some to have lysosomal activity.

References

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