Phototrophy

from Wikipedia, the free encyclopedia
The red alga Stylonema alsidii is phototrophic.

Phototrophy or photrophy (from ancient Greek φῶς phos = the light + τροφή trophé = the diet) describes the use of light as an energy source by living beings . The light is needed to synthesize the energy-rich chemical substance adenosine triphosphate (ATP) as an energy carrier and short-term energy store. With this ATP synthesis, living beings convert light energy into chemical energy.

Only certain organisms can use light energy directly for their metabolism. They are called phototrophic organisms or phototrophs . Phototrophy is common among prokaryotes (living things with cells without a nucleus) as well as among eukaryotes (living things with cells with a nucleus).

The phototrophic prokaryotes are metabolically divided into two groups. The first group uses light energy with the help of chlorophyll pigments ( chlorophylls or bacteriochlorophylls ). The second group is limited to a few archaea . They use light energy with the help of the rhodopsin pigments bacteriorhodopsin , proteorhodopsin or xanthorhodopsin . These are not structurally similar to chlorophyll pigments. In contrast, the phototrophic eukaryotes do not show any such metabolic physiological diversity.

Phototrophy should not be confused with phototropy (color change in crystalline substances and glasses due to exposure to light).

Forms of phototrophy

To perform phototrophy, organisms need very specific dyes ( pigments ). These special pigments are located in biomembranes . There they absorb light and make the radiation energy contained in it usable. So far, two different classes of such pigments have been discovered in phototrophic organisms: chlorophylls (chlorophylls, bacteriochlorophylls) and rhodopsins (bacteriorhodopsin, proteorhodopsin, xanthorhodopsin).

Phototrophy with chlorophylls: light-dependent reaction of photosynthesis

Chlorophyll and electron transport chain
The green alga Scenedesmus dimorphus becomes phototrophic through photosynthesis.

During photosynthesis , chlorophyll (or bacteriochlorophyll) is converted from its basic chemical state into an energetic (“excited”) state by the energy of light. In an excited state, a chlorophyll molecule easily gives off an energetic electron.

The electron is passed on via certain other molecules ( electron transport chain ), which are also located in the biomembrane. In the course of electron transport, hydrogen ions ( protons , H + ) are scooped from one side of the biomembrane to the other. That is why their concentration increases on one side of the membrane and at the same time decreases on the other side. This creates a high H + concentration gradient between the two sides of the membrane ( proton gradient ).

The proton gradient is used to build up ATP: the ATP synthase is embedded in the biomembrane . This is an enzyme , the synthesis of ATP from adenosine diphosphate (ADP) and phosphate (P i ) catalyzes . A channel runs within the ATP synthase that connects both sides of the membrane. Protons flow through the channel along their concentration gradient (→ diffusion ). The kinetic energy of the hydrogen nuclei flowing through is used by the ATP synthase for the synthesis of ATP, i.e. converted into chemical energy (→ chemiosmosis ).

Light energy is used to build up a proton gradient. The proton gradient is used to synthesize ATP. This phototrophic process is part of the so-called light - dependent reaction of photosynthesis.

The connection of P i to other substances is called phosphorylation . During the light-dependent reaction, ADP is phosphorylated to ATP with the help of light. Hence the process is called photophosphorylation .

A distinction is made between different forms of photosynthesis. During oxygenic photosynthesis , water molecules are split. The splitting of the water molecules ( photolysis ) also requires light and chlorophyll. The electron supply for the electron transport chain is obtained from the split water molecules. In addition, oxygen is released. In anoxygenic photosynthesis , other organic or inorganic substances are used for the supply of electrons and not water. No light is required for this and no oxygen is released.

inorganic electron donors in photosynthesis
Electron donation (at) or Photosynthesis form Occurrence
Iron-II-ions (Fe 2+ ) anoxygenic photosynthesis Purple bacteria
Nitrite (NO 2 - ) anoxygenic photosynthesis Purple bacteria
elemental sulfur (S 0 ) anoxygenic photosynthesis Purple bacteria
Hydrogen sulfide (H 2 S) anoxygenic photosynthesis green non-sulfur bacteria, green sulfur bacteria, purple bacteria
Thiosulfate (S 2 O 3 2− ) anoxygenic photosynthesis Purple bacteria
Water (H 2 O) oxygenic photosynthesis Cyanobacteria, phototrophic eukaryotes
Hydrogen (H 2 ) anoxygenic photosynthesis green non-sulfur bacteria

Phototrophy with rhodopsins

Haloarchaea become phototrophic with bacteriorhodopsin.
Proton pump rhodopsin

The phototrophic ATP synthesis (photophosphorylation) with the help of bacteriorhodopsin , proteorhodopsin or xanthorhodopsin also proceeds chemiosmotically . A rhodopsin consists of a protein that spans an entire biomembrane ( transmembrane protein ). Inside the protein is a molecule called retinal .

If light of a certain wavelength hits the retinal, the molecule changes its shape. In the process, a proton is released to the outside of the biomembrane. A new hydrogen nucleus is then fed to the retinal from the inside of the membrane. With the new proton, the molecule falls back into its original shape - until it is hit by light again, changes its shape again and releases a proton to the outside of the membrane once more. In this way, a rhodopsin-based phototrophy creates a high H + concentration gradient between both sides of the membrane (proton gradient).
The proton gradient is broken down by the fact that protons flow back through the channel of an ATP synthase along their concentration gradient to the inside of the membrane. The kinetic energy of the hydrogen nuclei flowing through is used for the synthesis of ATP (→ function of bacteriorhodopsin ).

Phototrophic organisms

Phototrophic prokaryotes

Different prokaryotes have evolved different forms of phototrophy. On the one hand, ATP synthesis developed with the help of the pigments bacteriorhodopsin, proteorhodopsin or xanthorhodopsin. On the other hand, phototrophy evolved independently with the help of chlorophyll pigments, photosynthesis. For many of the phototrophic prokaryotes, phototrophy is not the only option for energy metabolism. Especially in the dark, you may be able to use different ways of providing chemotrophic energy.

Phototrophic prokaryotes
Prokaryotes Substance and energy change Phototrophy form
Cyanobacteria Photolithoautotrophy (photohydroautotrophy) oxygenic photosynthesis
green non-sulfur bacteria Photoorganoheterotrophy or photolithoautotrophy Type II anoxygenic photosynthesis
green sulfur bacteria Photolithoautotrophy or photoorganoheterotrophy Type I anoxygenic photosynthesis , even at hydrothermal springs in the deep sea
Haloarchaea Photoorganoheterotrophy Phototrophy with bacteriorhodopsin
Heliobacteria Photoorganoheterotrophy Type I anoxygenic photosynthesis
Non-sulfur purple bacteria Photolithoautotrophy or photoorganoheterotrophy Type II anoxygenic photosynthesis
Thermoplasmates Photoorganoheterotrophy Phototrophy with proteorhodopsin
α-proteobacteria Photoorganoheterotrophy anoxygenic photosynthesis of type II or phototrophy with proteorhodopsin
Fulvimarina pelagi

(an α-proteobacterium)

Photoorganoheterotrophy Type II anoxygenic photosynthesis and phototrophy with xanthorhodopsin
β-proteobacteria Photoorganoheterotrophy Type II anoxygenic photosynthesis
γ-proteobacteria Photoorganoheterotrophy anoxygenic photosynthesis of type II or phototrophy with proteorhodopsin
Flavobacteria Photoorganoheterotrophy Phototrophy with proteorhodopsin
Salinibacter ruber

(an extremely halophilic bacterium )

Photoorganoheterotrophy Phototrophy with xanthorhodopsin
Sulfur purple bacteria Photolithoautotrophy Type II anoxygenic photosynthesis

Phototrophic eukaryotes

Originally, eukaryotes were not phototrophic. However, some eukaryotes acquired the ability to phototrophy by forming communities for mutual benefit ( mutualisms ) with phototrophic organisms. Such mutualisms arose several times, at different times and independently of one another. Plastids represent the most advanced form of phototrophic mutualism (→ endosymbiotic theory ).

Most of the phototrophic eukaryotes practice some form of photolithoautotrophy, which is sometimes called separately as photohydroautotrophy. This means that they synthesize carbohydrates exclusively with light, water and carbon dioxide. These phototrophic eukaryotes carry out oxygenic photosynthesis with the help of their mutualistic partners. Phototrophy is not the only source of nutrients for many phototrophic eukaryotes. They can also feed themselves chemoorganoheterotrophically by supplying themselves with nutrients by completely or partially eating other organisms. Such life forms, both autotrophically and heterotrophically, practice mixotrophy . It was only recognized a few years ago that many organisms that were previously classified as autotrophic or heterotrophic are in reality mixotrophic.

Algae and land plants

The most famous phototrophic eukaryotes harbor phototrophic plastids, the chloroplasts, in their cells. The chloroplast organelle emerged only once and that about 1.6 billion years ago. At that time it was possible to keep a cyanobacterium permanently inside the eukaryotic cell. So-called primary chloroplasts were formed. The primary chloroplasts of red algae and glaucophytes, which are presumably homologous to the other chloroplasts, are (mainly for historical reasons) called rhodoplasts (in red algae) and as muroplasts (in the past also: cyanelles) in glaucophytes.

Other eukaryotes later ingested such eukaryotes together with their primary chloroplasts (or alternatively: rhodoplasts). The ingested eukaryotes were gradually reduced until almost only their primary chloroplasts were left. In this way, secondary, complex chloroplasts were formed . Unlike the primary chloroplasts, these are then surrounded by four cell membranes instead of two. The inner two membranes go back to the chloroplast itself, the third to an enclosing vacuole and the fourth to the original cell membrane of the enclosed organism that contained the chloroplasts. In many cases, no further remnants of this organism are morphologically recognizable, but genes that originate from the host can still be detected, subsequently integrated into the host's core genome.

The uptake of chloroplasts was particularly varied in the Dinophyceae or armored flagellants. In this group, chloroplasts were apparently obtained multiple times and independently of one another. Most armored flagellants have secondary chloroplasts from red algae, but others have chloroplasts derived from recruited green algae. A small group of Dinophyceae even has tertiary chloroplasts. They come from ingested diatoms , gullet or calcareous algae , which in turn possessed secondary chloroplasts.

When the phototrophic eukaryotic cells divide , their chloroplasts multiply in them in parallel. Eukaryotes with such chloroplasts are grouped under the name algae . Most algae are microalgae . They remain microscopic, often live as protozoa or form cell colonies or coenobia with a very limited number of cells. Few groups of algae have developed multicellular forms. Such macroalgae are found exclusively among the Rhodophyta (red algae), the Phaeophyta (brown algae) and the Chloroplastida (green algae and the like).

A group of algae managed to colonize the land permanently; today they form land plants (next to them, often unicellular, microalgae occur as colonies or coatings on land). They belonged to the Streptophytina , which belong to the Charophyta , which in turn are assigned to the Chloroplastida. The chloroplasts of all chloroplastida appear green. The oldest fossils of green land plants are 475 million years old. These are spores of plants that are likely to have belonged to the liverworts . Today's land plants can be divided into mosses and tracheophyta ( vascular plants ).

Algae i. w. S. and land plants
large eukaryotic group phototrophic mutualist
Chlorarachniophyta ( green pseudopods ) complex chloroplasts
Chloroplastida : Chlorophyta ( green algae ), Embryophyta ( land plants ) u. a. Chloroplasts
Chrysophyta ( golden algae ): Bacillariophyceae ( diatoms ), Chrysophyceae ( golden algae ), Xanthophyceae ( yellow-green algae ) complex rhodoplasts
Cryptophyta ( gullet ) complex rhodoplasts
Dinophyta ( armored flagellant ) complex rhodoplasts
Euglenophyta ( flagella ) complex chloroplasts
Glaucophyta Muroplasts (formerly called cyanelles)
Haptophyta ( calcareous algae ) complex rhodoplasts
Phaeophyta ( brown algae ) Phaeoplasts (complex rhodoplasts)
Rhodophyta ( red algae ) Rhodoplasts

Phototrophic protists without chloroplasts

In addition to the phototrophic protists with chloroplasts, there are many other phototrophic unicellular organisms that do not have chloroplasts. These protists acquire phototrophy in other ways. It often happens by ingesting other phototrophic protozoa - cyanobacteria or zoochlorella.

In addition to photosynthesis based on chlorophyll, primary energy production (not based on endosymbionts) using the rhodopsin pigment proteorhodopsin through DNA sequencing was detected not only in numerous autotrophic, but also a large number of heterotrophic protists, which are actually forms with a mixed diet (mixotrophic ) would have to be classified, the ecological consequences of which have so far been poorly understood. This ability has also passed to autotrophic and, to a few heterotrophic, eukaryotes via symbiogenesis. The (heterotrophic) dinoflagellate Oxyrrhis marina even has numerous rhodopsins from two different families: In addition to the sensory rhodopsins widespread in eukaryotes, proteorhodopsins have also been detected, which drive a proton pump using sunlight and thus contribute to energy production; It has been shown experimentally that the species survives periods of starvation longer when exposed than in darkness. Corresponding proteorhodopsins were later also detected in other autotrophic and heterotrophic dinoflagellates.

Phototrophic protists without chloroplasts (small selection)
Protist large eukaryotic group particular phototrophic mutualist

or other form of phototrophy

Auranticordis quadriverberis Cercozoa Cyanobacteria
Large foraminifera Foraminifera ( chamberlings ) Zoochlorella or zooxanthellae
Hatena arenicola Cryptophyta ( gullet ) Nephroselmis (a green algae)
Oxyrrhis marina Dinozoa Phototrophy with proteorhodopsin
Paramecium bursaria Ciliophora ( eyelash animal ) Zoochlorella
Paulinella chromatophora Cercozoa Cyanobacteria ( Prochlorococcus or Synechococcus )
Stentor polymorphus Ciliophora (eyelash animal) Zoochlorella

Phototrophic opisthoconta

Not only algae, land plants and certain protists do phototrophy. Even among the opisthokonta - that is, among fungi and animals - some groups were able to switch to the phototrophic way of life. Phototrophy was achieved in very different ways.

  • Phototrophy with photobionts : Lichenes ( lichens ) are mutualisms between fungi and phototrophic protozoa, which are collectively called photobionts . Many photobionts are single- or few-celled chlorophyta (green algae). The remaining photobionts come from the group of cyanobacteria.
  • Green algae of the genus Chlorella often live as phototrophic mutualists within other cells.
    Phototrophy with zoochlorella : Different animals evolved mutualisms between themselves and endosymbiotic green algae of the genus Chlorella ( zoochlorella ). Better known representatives are the green freshwater sponges Ephydatia fluviatilis and Spongilla lacustris . The cnidaria ( cnidarians ) include the green hydra ( Hydra viridissima ), which stores chlorella in vacuoles inside their cells. The cnidaria also include the green giant anemones Anthopleura elegantissima and Anthopleura xanthogrammica . The green giant anemones can not only harbor zoochlorella, but also zooxanthellae.
  • Phototrophy with other green algae : In addition to the frequently occurring zoochlorella, other single-celled chlorophyta (green algae) also entered into phototrophic mutualisms with animals. The green alga Tetraselmis convolutae lives between the cells of the green acoelomorph Symsagittifera roscoffensis . And Oophila amblystomatis lives in eggs and embryos of Ambystoma maculatum ( spotted transverse newt ). This newt is the only known example of a phototrophic vertebrate.
  • Phototrophy with zooxanthellae : Even more often than zoochlorella, zooxanthellae are used as phototrophic mutualists by animals. The most widespread are zooxanthellae of the genus Symbiodinium , which belong to the armored flagellants . Many cnidarians harbor zooxanthellae, especially the various corals . But zooxanthellae are also found in many sea anemones, for example, in the wake Rose ( Anemonia sulcata ), in the sun rose ( Cereus pedunculatus ) and - in addition zoochlorella - in the green giant anemones Anthopleura elegantissima and Anthopleura xanthogrammica . Zooxanthellae are still found in some jellyfish . For example in the umbrella jellyfish Mastigias papua and Linuche unguiculata and in the mangrove jellyfish Cassiopea xamachana . Outside of the cnidarians, only a few cases of animal mutualisms with zooxanthellae have so far been found. At least they occur in the Acoelomorpha genus Waminoa and in Tridacnidae, the giant clams .
  • Phototrophy with kleptoplastids : representatives of the sacoglossa ( gastropod snails ) rob chloroplasts from green algae. You are doing kleptoplasty . To do this, they eat cells from certain green algae. In the intestine, the chloroplasts are removed from the eaten cell contents. They are then transported through the screw body. Finally, the chloroplasts are stored in vacuoles in skin cells. These chloroplasts, which were originally stolen by the eaten green algae, are called klepto (chloro) plast (id) en . The best-known phototrophic representatives of the gastropod snails are the green velvet snail ( Elysia viridis ) and its close relative Elysia chlorotica .
  • Phototrophy with xanthopterin : The oriental hornet ( Vespa orientalis ) seems to have evolved a completely new way of using sunlight. To do this, she uses the yellow dye xanthopterin (in conjunction with the brown dye melanin ) in the cuticle of her exoskeleton in order to produce electricity in the same way as a solar cell. The biological purpose of the structure is not yet known. It may be used to generate chemical energy that helps the wasp dig underground nest cavities.

See also

literature

  • Donat-Peter Häder (Ed.): Photosynthesis . Stuttgart / New York 1999, ISBN 3-13-115021-1 .
  • MT Madigan, JM Martinko: Brock Microbiology . Munich 2006, ISBN 3-8273-7187-2 .
  • U. Sonnewald: Physiology. In: Strasburger textbook of botany. Heidelberg 2008, ISBN 978-3-8274-1455-7 .
  • J. Overmann, F. Garcia-Pichel: The Phototrophic Way of Life. In: The Prokaryotes, Vol. 2: Ecophysiology and Biochemistry. New York 2006, ISBN 0-387-30742-7 .

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