Chromoplast

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The coloration of the petals and sepals on the bee orchid is controlled by chromoplasts. Ophrys apifera flower1.jpg
The coloration of the petals and sepals on the bee orchid is controlled by chromoplasts.

Chromoplasts are plastids, heterogeneous organelles responsible for pigment synthesis and storage in specific photosynthetic eukaryotes. [1] It is thought that like all other plastids including chloroplasts and leucoplasts they are descended from symbiotic prokaryotes. [2]

Contents

Function

Chromoplasts are found in fruits, flowers, roots, and stressed and aging leaves, and are responsible for their distinctive colors. This is always associated with a massive increase in the accumulation of carotenoid pigments. The conversion of chloroplasts to chromoplasts in ripening is a classic example.

They are generally found in mature tissues and are derived from preexisting mature plastids. Fruits and flowers are the most common structures for the biosynthesis of carotenoids, although other reactions occur there as well including the synthesis of sugars, starches, lipids, aromatic compounds, vitamins, and hormones. [3] The DNA in chloroplasts and chromoplasts is identical. [2] One subtle difference in DNA was found after a liquid chromatography analysis of tomato chromoplasts was conducted, revealing increased cytosine methylation. [3]

Chromoplasts synthesize and store pigments such as orange carotene, yellow xanthophylls, and various other red pigments. As such, their color varies depending on what pigment they contain. The main evolutionary purpose of chromoplasts is probably to attract pollinators or eaters of colored fruits, which help disperse seeds. However, they are also found in roots such as carrots and sweet potatoes. They allow the accumulation of large quantities of water-insoluble compounds in otherwise watery parts of plants.

When leaves change color in the autumn, it is due to the loss of green chlorophyll, which unmasks preexisting carotenoids. In this case, relatively little new carotenoid is produced—the change in plastid pigments associated with leaf senescence is somewhat different from the active conversion to chromoplasts observed in fruit and flowers.

There are some species of flowering plants that contain little to no carotenoids. In such cases, there are plastids present within the petals that closely resemble chromoplasts and are sometimes visually indistinguishable. Anthocyanins and flavonoids located in the cell vacuoles are responsible for other colors of pigment. [1]

The term "chromoplast" is occasionally used to include any plastid that has pigment, mostly to emphasize the difference between them and the various types of leucoplasts, plastids that have no pigments. In this sense, chloroplasts are a specific type of chromoplast. Still, "chromoplast" is more often used to denote plastids with pigments other than chlorophyll.

Structure and classification

Using a light microscope chromoplasts can be differentiated and are classified into four main types. The first type is composed of proteic stroma with granules. The second is composed of protein crystals and amorphous pigment granules. The third type is composed of protein and pigment crystals. The fourth type is a chromoplast which only contains crystals. An electron microscope reveals even more, allowing for the identification of substructures such as globules, crystals, membranes, fibrils and tubules. The substructures found in chromoplasts are not found in the mature plastid that it divided from. [2]

The presence, frequency and identification of substructures using an electron microscope has led to further classification, dividing chromoplasts into five main categories: Globular chromoplasts, crystalline chromoplasts, fibrillar chromoplasts, tubular chromoplasts and membranous chromoplasts. [2] It has also been found that different types of chromoplasts can coexist in the same organ. [3] Some examples of plants in the various categories include mangoes, which have globular chromoplasts, and carrots which have crystalline chromoplasts. [4]

Although some chromoplasts are easily categorized, others have characteristics from multiple categories that make them hard to place. Tomatoes accumulate carotenoids, mainly lycopene crystalloids in membrane-shaped structures, which could place them in either the crystalline or membranous category. [3]

Evolution

Plastids lining which pollinators visit a flower, as specific colors attract specific pollinators. White flowers tend to attract beetles, bees are most often attracted to violet and blue flowers, and butterflies are often attracted to warmer colors like yellows and oranges. [5]

Research

Chromoplasts are not widely studied and are rarely the main focus of scientific research. They often play a role in research on the tomato plant ( Solanum lycopersicum ). Lycopene is responsible for the red color of a ripe fruit in the cultivated tomato, while the yellow color of the flowers is due to xanthophylls violaxanthin and neoxanthin. [6]

Carotenoid biosynthesis occurs in both chromoplasts and chloroplasts. In the chromoplasts of tomato flowers, carotenoid synthesis is regulated by the genes Psyl, Pds, Lcy-b, and Cyc-b. These genes, in addition to others, are responsible for the formation of carotenoids in organs and structures. For example, the Lcy-e gene is highly expressed in leaves, which results in the production of the carotenoid lutein. [6]

White flowers are caused by a recessive allele in tomato plants. They are less desirable in cultivated crops because they have a lower pollination rate. In one study, it was found that chromoplasts are still present in white flowers. The lack of yellow pigment in their petals and anthers is due to a mutation in the CrtR-b2 gene which disrupts the carotenoid biosynthesis pathway. [6]

The entire process of chromoplast formation is not yet completely understood on the molecular level. However, electron microscopy has revealed part of the transformation from chloroplast to chromoplast. The transformation starts with remodeling of the internal membrane system with the lysis of the intergranal thylakoids and the grana. New membrane systems form in organized membrane complexes called thylakoid plexus. The new membranes are the site of the formation of carotenoid crystals. These newly synthesized membranes do not come from the thylakoids, but rather from vesicles generated from the inner membrane of the plastid. The most obvious biochemical change would be the downregulation of photosynthetic gene expression which results in the loss of chlorophyll and stops photosynthetic activity. [3]

In oranges, the synthesis of carotenoids and the disappearance of chlorophyll causes the color of the fruit to change from green to yellow. The orange color is often added artificially—light yellow-orange is the natural color created by the actual chromoplasts. [7]

Valencia oranges Citris sinensis L are a cultivated orange grown extensively in the state of Florida. In the winter, Valencia oranges reach their optimum orange-rind color while reverting to a green color in the spring and summer. While it was originally thought that chromoplasts were the final stage of plastid development, in 1966 it was proved that chromoplasts can revert to chloroplasts, which causes the oranges to turn back to green. [7]

Compare plastids

Plastids types.svg

Related Research Articles

<span class="mw-page-title-main">Chloroplast</span> Plant organelle that conducts photosynthesis

A chloroplast is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in the energy-storage molecules ATP and NADPH while freeing oxygen from water in the cells. The ATP and NADPH is then used to make organic molecules from carbon dioxide in a process known as the Calvin cycle. Chloroplasts carry out a number of other functions, including fatty acid synthesis, amino acid synthesis, and the immune response in plants. The number of chloroplasts per cell varies from one, in unicellular algae, up to 100 in plants like Arabidopsis and wheat.

<span class="mw-page-title-main">Chlorophyll</span> Green pigments found in plants, algae and bacteria

Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words χλωρός, khloros and φύλλον, phyllon ("leaf"). Chlorophyll allow plants to absorb energy from light.

<span class="mw-page-title-main">Carotene</span> Class of compounds

The term carotene (also carotin, from the Latin carota, "carrot") is used for many related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but in general cannot be made by animals (with the exception of some aphids and spider mites which acquired the synthesizing genes from fungi). Carotenes are photosynthetic pigments important for photosynthesis. Carotenes contain no oxygen atoms. They absorb ultraviolet, violet, and blue light and scatter orange or red light, and (in low concentrations) yellow light.

<span class="mw-page-title-main">Lycopene</span> Carotenoid pigment

Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and vegetables.

<span class="mw-page-title-main">Photosynthesis</span> Biological process to convert light into chemical energy

Photosynthesis is a biological process used by many cellular organisms to convert light energy into chemical energy, which is stored in organic compounds that can later be metabolized through cellular respiration to fuel the organism's activities. The term usually refers to oxygenic photosynthesis, where oxygen is produced as a byproduct, and some of the chemical energy produced is stored in carbohydrate molecules such as sugars, starch and cellulose, which are synthesized from endergonic reaction of carbon dioxide with water. Most plants, algae and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the biological energy necessary for complex life on Earth.

<span class="mw-page-title-main">Photosynthetic pigment</span> Type of molecule

A photosynthetic pigment is a pigment that is present in chloroplasts or photosynthetic bacteria and captures the light energy necessary for photosynthesis.

<span class="mw-page-title-main">Plastid</span> Plant cell organelles that perform photosynthesis and store starch

The plastid is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyanobacteria. Examples include chloroplasts, chromoplasts, and leucoplasts.

<span class="mw-page-title-main">Thylakoid</span> Membrane enclosed compartments in chloroplasts and cyanobacteria

Thylakoids are membrane-bound compartments inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thylakoids frequently form stacks of disks referred to as grana. Grana are connected by intergranal or stromal thylakoids, which join granum stacks together as a single functional compartment.

<span class="mw-page-title-main">Carotenoid</span> Class of chemical compounds; yellow, orange or red plant pigments

Carotenoids are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, canaries, flamingos, salmon, lobster, shrimp, and daffodils. Over 1,100 identified carotenoids can be further categorized into two classes – xanthophylls and carotenes.

<span class="mw-page-title-main">Chloroplast membrane</span>

Chloroplasts contain several important membranes, vital for their function. Like mitochondria, chloroplasts have a double-membrane envelope, called the chloroplast envelope, but unlike mitochondria, chloroplasts also have internal membrane structures called thylakoids. Furthermore, one or two additional membranes may enclose chloroplasts in organisms that underwent secondary endosymbiosis, such as the euglenids and chlorarachniophytes.

<span class="mw-page-title-main">Xanthophyll</span> Chemical compounds subclass

Xanthophylls are yellow pigments that occur widely in nature and form one of two major divisions of the carotenoid group; the other division is formed by the carotenes. The name is from Greek: xanthos (ξανθός), meaning "yellow", and phyllon (φύλλον), meaning "leaf"), due to their formation of the yellow band seen in early chromatography of leaf pigments.

<span class="mw-page-title-main">Photosystem</span> Structural units of protein involved in photosynthesis

Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.

<span class="mw-page-title-main">Etioplast</span>

Etioplasts are an intermediate type of plastid that develop from proplastids that have not been exposed to light, and convert into chloroplasts upon exposure to light. They are usually found in stem and leaf tissue of flowering plants (Angiosperms) grown either in complete darkness, or in extremely low-light conditions.

<span class="mw-page-title-main">Biological pigment</span> Substances produced by living organisms

Biological pigments, also known simply as pigments or biochromes, are substances produced by living organisms that have a color resulting from selective color absorption. Biological pigments include plant pigments and flower pigments. Many biological structures, such as skin, eyes, feathers, fur and hair contain pigments such as melanin in specialized cells called chromatophores. In some species, pigments accrue over very long periods during an individual's lifespan.

<span class="mw-page-title-main">Proteinoplast</span> Organelles in plant cells

Proteinoplasts are specialized organelles found only in plant cells. Proteinoplasts belong to a broad category of organelles known as plastids. Plastids are specialized double-membrane organelles found in plant cells. Plastids perform a variety of functions such as metabolism of energy, and biological reactions. There are multiple types of plastids recognized including Leucoplasts, Chromoplasts, and Chloroplasts. Plastids are broken up into different categories based on characteristics such as size, function and physical traits. Chromoplasts help to synthesize and store large amounts of carotenoids. Chloroplasts are photosynthesizing structures that help to make light energy for the plant. Leucoplasts are a colorless type of plastid which means that no photosynthesis occurs here. The colorless pigmentation of the leucoplast is due to not containing the structural components of thylakoids unlike what is found in chloroplasts and chromoplasts that gives them their pigmentation. From leucoplasts stems the subtype, proteinoplasts, which contain proteins for storage. They contain crystalline bodies of protein and can be the sites of enzyme activity involving those proteins. Proteinoplasts are found in many seeds, such as brazil nuts, peanuts and pulses. Although all plastids contain high concentrations of protein, proteinoplasts were identified in the 1960s and 1970s as having large protein inclusions that are visible with both light microscopes and electron microscopes. Other subtypes of Leucoplasts include amyloplast, and elaioplasts. Amyloplasts help to store and synthesize starch molecules found in plants, while elaioplasts synthesize and store lipids in plant cells.

<span class="mw-page-title-main">Diadinoxanthin</span> Chemical compound

Diadinoxanthin is a pigment found in phytoplankton. It has the formula C40H54O3. It gives rise to the xanthophylls diatoxanthin and dinoxanthin.

Antheraxanthin is a bright yellow accessory pigment found in many organisms that perform photosynthesis. It is a xanthophyll cycle pigment, an oil-soluble alcohol within the xanthophyll subgroup of carotenoids. Antheraxanthin is both a component in and product of the cellular photoprotection mechanisms in photosynthetic green algae, red algae, euglenoids, and plants.

<span class="mw-page-title-main">15-Cis-phytoene desaturase</span> Class of enzymes

15-cis-phytoene desaturases, are enzymes involved in the carotenoid biosynthesis in plants and cyanobacteria. Phytoene desaturases are membrane-bound enzymes localized in plastids and introduce two double bonds into their colorless substrate phytoene by dehydrogenation and isomerize two additional double bonds. This reaction starts a biochemical pathway involving three further enzymes called the poly-cis pathway and leads to the red colored lycopene. The homologous phytoene desaturase found in bacteria and fungi (CrtI) converts phytoene directly to lycopene by an all-trans pathway.

Lycopene β-cyclase is an enzyme with systematic name carotenoid beta-end group lyase (decyclizing). This enzyme catalyses the following chemical reaction

Myxoxanthophyll is a carotenoid glycoside pigment present in the photosynthetic apparatus of cyanobacteria. It is named after the word "Myxophyceae", a former term for cyanobacteria. As a monocyclic xanthophyll, it has a yellowish color. It is required for normal cell wall structure and thylakoid organization in the cyanobacterium Synechocystis. The pigment is unusual because it is glycosylated on the 2'-OH rather than the 1'-OH position of the molecule. Myxoxanthophyll was first isolated from Oscillatoria rubenscens in 1936.

References

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