Evolution of photosynthesis

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The evolution of photosynthesis refers to the origin and subsequent evolution of photosynthesis, the process by which light energy is used to assemble sugars from carbon dioxide and a hydrogen and electron source such as water. The process of photosynthesis was discovered by Jan Ingenhousz, a Dutch-born British physician and scientist, first publishing about it in 1779. [1]

Contents

The first photosynthetic organisms probably evolved early in the evolutionary history of life and most likely used reducing agents such as hydrogen rather than water. [2] There are three major metabolic pathways by which photosynthesis is carried out: C3 photosynthesis, C4 photosynthesis, and CAM photosynthesis. C3 photosynthesis is the oldest and most common form. A C3 plant uses the Calvin cycle for the initial steps that incorporate CO2 into organic material. A C4 plant prefaces the Calvin cycle with reactions that incorporate CO2 into four-carbon compounds. A CAM plant uses crassulacean acid metabolism, an adaptation for photosynthesis in arid conditions. C4 and CAM plants have special adaptations that save water. [3]

Origin

Available evidence from geobiological studies of Archean (>2500 Ma) sedimentary rocks indicates that life existed 3500 Ma. Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old, [4] [5] consistent with recent studies of photosynthesis. [6] [7] Early photosynthetic systems, such as those from green and purple sulfur and green and purple nonsulfur bacteria, are thought to have been anoxygenic, using various molecules as electron donors. Green and purple sulfur bacteria are thought to have used hydrogen and hydrogen sulfide as electron and hydrogen donors. Green nonsulfur bacteria used various amino and other organic acids. Purple nonsulfur bacteria used a variety of nonspecific organic and inorganic molecules. [8] It is suggested that photosynthesis likely originated at low-wavelength geothermal light from acidic hydrothermal vents, Zn-tetrapyrroles were the first photochemically active pigments, the photosynthetic organisms were anaerobic and relied on H2S without relying on H2 emitted by alkaline hydrothermal vents. The divergence of anoxygenic photosynthetic organisms at the photic zone could have led to the ability to strip electrons from H2S more efficiently under ultraviolet radiation. There is geochemical evidence that suggests that anaerobic photosynthesis emerged 3.3 to 3.5 billion years ago. The organisms later developed a Chlorophyll F synthase. They could have also stripped electrons from soluble metal ions although it is unknown. [9]

The first oxygenic photosynthetic organisms are proposed to be H2S-dependent. [9] It is also suggested photosynthesis originated under sunlight, using H2S emitted by volcanoes and hydrothermal vents which ended the need for scarce H2 emitted by alkaline hydrothermal vents. [10] Oxygenic photosynthesis uses water as an electron donor, which is oxidized to molecular oxygen (O
2) in the photosynthetic reaction center. The biochemical capacity for oxygenic photosynthesis evolved in a common ancestor of extant cyanobacteria. [11] The first appearance of free oxygen in the atmosphere is sometimes referred to as the oxygen catastrophe. The geological record indicates that this transforming event took place during the Paleoproterozoic era at least 2450–2320 million years ago (Ma), and, it is speculated, much earlier. [12] [13] A clear paleontological window on cyanobacterial evolution opened about 2000 Ma, revealing an already-diverse biota of blue-greens. Cyanobacteria remained principal primary producers throughout the Proterozoic Eon (2500–543 Ma), in part because the redox structure of the oceans favored photoautotrophs capable of nitrogen fixation. [14] Green algae joined blue-greens as major primary producers on continental shelves near the end of the Proterozoic, but only with the Mesozoic (251–65 Ma) radiations of dinoflagellates, coccolithophorids, and diatoms did primary production in marine shelf waters take modern form. Cyanobacteria remain critical to marine ecosystems as primary producers in oceanic gyres, as agents of biological nitrogen fixation, and, in modified form, as the plastids of marine algae. [15] Modern photosynthesis in plants and most photosynthetic prokaryotes is oxygenic.

Timeline of photosynthesis on Earth

4.6 billion years agoEarth forms
3.4 billion years agoFirst photosynthetic bacteria appear
2.7 billion years agoCyanobacteria become the first oxygen producers
2.4 – 2.3 billion years agoEarliest evidence (from rocks) that oxygen was in the atmosphere
1.2 billion years agoRed and brown algae become structurally more complex than bacteria
0.75 billion years agoGreen algae outperform red and brown algae in the strong light of shallow water
0.475 billion years agoFirst land plants – mosses and liverworts
0.423 billion years agoVascular plants evolve

Source: [16]

Symbiosis and the origin of chloroplasts

Plant cells with visible chloroplasts (from a moss, Plagiomnium affine) Plagiomnium affine laminazellen.jpeg
Plant cells with visible chloroplasts (from a moss, Plagiomnium affine )

Several groups of animals have formed symbiotic relationships with photosynthetic algae. These are most common in corals, sponges and sea anemones. It is presumed that this is due to the particularly simple body plans and large surface areas of these animals compared to their volumes. [17] In addition, a few marine mollusks Elysia viridis and Elysia chlorotica also maintain a symbiotic relationship with chloroplasts they capture from the algae in their diet and then store in their bodies. This allows the mollusks to survive solely by photosynthesis for several months at a time. [18] [19] Some of the genes from the plant cell nucleus have even been transferred to the slugs, so that the chloroplasts can be supplied with proteins that they need to survive. [20]

An even closer form of symbiosis may explain the origin of chloroplasts. Chloroplasts have many similarities with photosynthetic bacteria, including a circular chromosome, prokaryotic-type ribosomes, and similar proteins in the photosynthetic reaction center. [21] [22] The endosymbiotic theory suggests that photosynthetic bacteria were acquired (by endocytosis) by early eukaryotic cells to form the first plant cells. Therefore, chloroplasts may be photosynthetic bacteria that adapted to life inside plant cells. Like mitochondria, chloroplasts still possess their own DNA, separate from the nuclear DNA of their plant host cells and the genes in this chloroplast DNA resemble those in cyanobacteria. [23] DNA in chloroplasts codes for redox proteins such as photosynthetic reaction centers. The CoRR Hypothesis proposes that this Co-location is required for Redox Regulation.

Evolution of photosynthetic pathways

The C4 carbon concentrating mechanism HatchSlackpathway2.svg
The C4 carbon concentrating mechanism

In its simplest form, photosynthesis is adding water to CO2 to produce sugars and oxygen, but a complex chemical pathway is involved, facilitated along the way by a range of enzymes and co-enzymes. The enzyme RuBisCO is responsible for "fixing" CO2 – that is, it attaches it to a carbon-based molecule to form a sugar, which can be used by the plant, releasing an oxygen molecule along the way. However, the enzyme is notoriously inefficient, and just as effectively will also fix oxygen instead of CO2 in a process called photorespiration. This is energetically costly as the plant has to use energy to turn the products of photorespiration back into a form that can react with CO2.[ citation needed ] [24]

Concentrating carbon

The C4 metabolic pathway is a valuable recent evolutionary innovation in plants, involving a complex set of adaptive changes to physiology and gene expression patterns. [25] About 7600 species of plants use C4 carbon fixation, which represents about 3% of all terrestrial species of plants. All these 7600 species are angiosperms.

C4 plants evolved carbon concentrating mechanisms. These work by increasing the concentration of CO2 around RuBisCO, thereby facilitating photosynthesis and decreasing photorespiration. The process of concentrating CO2 around RuBisCO requires more energy than allowing gases to diffuse, but under certain conditions – i.e. warm temperatures (>25 °C), low CO2 concentrations, or high oxygen concentrations – pays off in terms of the decreased loss of sugars through photorespiration.[ citation needed ]

One type of C4 metabolism employs a so-called Kranz anatomy. This transports CO2 through an outer mesophyll layer, via a range of organic molecules, to the central bundle sheath cells, where the CO2 is released. In this way, CO2 is concentrated near the site of RuBisCO operation. Because RuBisCO is operating in an environment with much more CO2 than it otherwise would be, it performs more efficiently.[ citation needed ] [26] In C4 photosynthesis, carbon is fixed by an enzyme called PEP carboxylase, which, like all enzymes involved in C4 photosynthesis, originated from non-photosynthetic ancestral enzymes. [27] [28]

A second mechanism, CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions. [29] [30] The most important benefit of CAM to the plant is the ability to leave most leaf stomata closed during the day. [31] This reduces water loss due to evapotranspiration. The stomata open at night to collect CO2, which is stored as the four-carbon acid malate, and then used during photosynthesis during the day. The pre-collected CO2 is concentrated around the enzyme RuBisCO, increasing photosynthetic efficiency. More CO2 is then harvested from the atmosphere when stomata open, during the cool, moist nights, reducing water loss.[ citation needed ]

CAM has evolved convergently many times. [32] It occurs in 16,000 species (about 7% of plants), belonging to over 300 genera and around 40 families, but this is thought to be a considerable underestimate. [33] It is found in quillworts (relatives of club mosses), in ferns, and in gymnosperms, but the great majority of plants using CAM are angiosperms (flowering plants).[ citation needed ]

Evolutionary record

These two pathways, with the same effect on RuBisCO, evolved a number of times independently – indeed, C4 alone arose 62 times in 18 different plant families. A number of 'pre-adaptations' seem to have paved the way for C4, leading to its clustering in certain clades: it has most frequently developed in plants that already had features such as extensive vascular bundle sheath tissue. [34] Whole-genome and individual gene duplication are also associated with C4 evolution. [35] Many potential evolutionary pathways resulting in the C4 phenotype are possible and have been characterised using Bayesian inference, [25] confirming that non-photosynthetic adaptations often provide evolutionary stepping stones for the further evolution of C4.

Crassulacean Acid Metabolism (CAM) is named after the family Crassulaceae, to which the jade plant belongs. Another example of a CAM plant is the pineapple. Crassula Ovata.jpg
Crassulacean Acid Metabolism (CAM) is named after the family Crassulaceae, to which the jade plant belongs. Another example of a CAM plant is the pineapple.

The C4 construction is most famously used by a subset of grasses, while CAM is employed by many succulents and cacti. The trait appears to have emerged during the Oligocene, around 25 to 32 million years ago; [36] however, they did not become ecologically significant until the Miocene, 6 to 7 million years ago. [37] Remarkably, some charcoalified fossils preserve tissue organised into the Kranz anatomy, with intact bundle sheath cells, [38] allowing the presence C4 metabolism to be identified without doubt at this time. Isotopic markers are used to deduce their distribution and significance.

C3 plants preferentially use the lighter of two isotopes of carbon in the atmosphere, 12C, which is more readily involved in the chemical pathways involved in its fixation. Because C4 metabolism involves a further chemical step, this effect is accentuated. Plant material can be analysed to deduce the ratio of the heavier 13C to 12C. This ratio is denoted δ13C. C3 plants are on average around 14‰ (parts per thousand) lighter than the atmospheric ratio, while C4 plants are about 28‰ lighter. The δ13C of CAM plants depends on the percentage of carbon fixed at night relative to what is fixed in the day, being closer to C3 plants if they fix most carbon in the day and closer to C4 plants if they fix all their carbon at night. [39]

It is troublesome procuring original fossil material in sufficient quantity to analyse the grass itself, but fortunately there is a good proxy: horses. Horses were globally widespread in the period of interest, and browsed almost exclusively on grasses. There's an old phrase in isotope palæontology, "you are what you eat (plus a little bit)" – this refers to the fact that organisms reflect the isotopic composition of whatever they eat, plus a small adjustment factor. There is a good record of horse teeth throughout the globe, and their δ13C has been measured. The record shows a sharp negative inflection around 6 to 7 million years ago, during the Messinian, and this is interpreted as the rise of C4 plants on a global scale. [37]

When is C4 an advantage?

While C4 enhances the efficiency of RuBisCO, the concentration of carbon is highly energy intensive. This means that C4 plants only have an advantage over C3 organisms in certain conditions: namely, high temperatures and low rainfall. C4 plants also need high levels of sunlight to thrive. [40] Models suggest that, without wildfires removing shade-casting trees and shrubs, there would be no space for C4 plants. [41] But, wildfires have occurred for 400 million years – why did C4 take so long to arise, and then appear independently so many times? The Carboniferous period (~ 300  million years ago) had notoriously high oxygen levels – almost enough to allow spontaneous combustion [42]  – and very low CO2, but there is no C4 isotopic signature to be found. And there doesn't seem to be a sudden trigger for the Miocene rise.[ citation needed ]

During the Miocene, the atmosphere and climate were relatively stable. If anything, CO2 increased gradually from 14 to 9 million years ago before settling down to concentrations similar to the Holocene. [43] This suggests that it did not have a key role in invoking C4 evolution. [36] Grasses themselves (the group which would give rise to the most occurrences of C4) had probably been around for 60 million years or more, so had had plenty of time to evolve C4, [44] [45] which, in any case, is present in a diverse range of groups and thus evolved independently. There is a strong signal of climate change in South Asia; [36] increasing aridity – hence increasing fire frequency and intensity – may have led to an increase in the importance of grasslands. [46] However, this is difficult to reconcile with the North American record. [36] It is possible that the signal is entirely biological, forced by the fire- and grazer- [47] driven acceleration of grass evolution – which, both by increasing weathering and incorporating more carbon into sediments, reduced atmospheric CO2 levels. [47] Finally, there is evidence that the onset of C4 from 9 to 7 million years ago is a biased signal, which only holds true for North America, from where most samples originate; emerging evidence suggests that grasslands evolved to a dominant state at least 15Ma earlier in South America.[ citation needed ]

See also

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">Photosynthesis</span> Biological process to convert light into chemical energy

Photosynthesis is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their activities. Photosynthetic organisms use intracellular organic compounds to store the chemical energy they produce in photosynthesis. Photosynthesis is usually used to refer to oxygenic photosynthesis, a form of photosynthesis where the photosynthetic processes produce oxygen as a byproduct and synthesize carbohydrate molecules like sugars, starches, glycogen, and cellulose to store the chemical energy. To use the chemical energy stored in these organic compounds, the organisms' cells metabolize the organic compounds through another process called cellular respiration. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and it supplies most of the biological energy necessary for complex life on Earth.

<span class="mw-page-title-main">RuBisCO</span> Key enzyme of photosynthesis involved in carbon fixation

Ribulose-1,5-bisphosphate carboxylase/oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme involved in light-independent part of photosynthesis, including the carbon fixation by which atmospheric carbon dioxide is converted by plants and other photosynthetic organisms to energy-rich molecules such as glucose. It emerged approximately four billion years ago in primordial metabolism prior to the presence of oxygen on Earth. It is probably the most abundant enzyme on Earth. In chemical terms, it catalyzes the carboxylation of ribulose-1,5-bisphosphate.

<span class="mw-page-title-main">Crassulacean acid metabolism</span> Metabolic process

Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions that allows a plant to photosynthesize during the day, but only exchange gases at night. In a plant using full CAM, the stomata in the leaves remain shut during the day to reduce evapotranspiration, but they open at night to collect carbon dioxide and allow it to diffuse into the mesophyll cells. The CO2 is stored as four-carbon malic acid in vacuoles at night, and then in the daytime, the malate is transported to chloroplasts where it is converted back to CO2, which is then used during photosynthesis. The pre-collected CO2 is concentrated around the enzyme RuBisCO, increasing photosynthetic efficiency. This mechanism of acid metabolism was first discovered in plants of the family Crassulaceae.

C<sub>4</sub> carbon fixation Photosynthetic process in some plants

C4 carbon fixation or the Hatch–Slack pathway is one of three known photosynthetic processes of carbon fixation in plants. It owes the names to the 1960s discovery by Marshall Davidson Hatch and Charles Roger Slack.

<span class="mw-page-title-main">Photorespiration</span> Process in plant metabolism

Photorespiration (also known as the oxidative photosynthetic carbon cycle or C2 cycle) refers to a process in plant metabolism where the enzyme RuBisCO oxygenates RuBP, wasting some of the energy produced by photosynthesis. The desired reaction is the addition of carbon dioxide to RuBP (carboxylation), a key step in the Calvin–Benson cycle, but approximately 25% of reactions by RuBisCO instead add oxygen to RuBP (oxygenation), creating a product that cannot be used within the Calvin–Benson cycle. This process lowers the efficiency of photosynthesis, potentially lowering photosynthetic output by 25% in C3 plants. Photorespiration involves a complex network of enzyme reactions that exchange metabolites between chloroplasts, leaf peroxisomes and mitochondria.

<span class="mw-page-title-main">Ribulose 1,5-bisphosphate</span> Chemical compound

Ribulose 1,5-bisphosphate (RuBP) is an organic substance that is involved in photosynthesis, notably as the principal CO2 acceptor in plants. It is a colourless anion, a double phosphate ester of the ketopentose called ribulose. Salts of RuBP can be isolated, but its crucial biological function happens in solution. RuBP occurs not only in plants but in all domains of life, including Archaea, Bacteria, and Eukarya.

<i>Chloroflexus aurantiacus</i> Species of bacterium

Chloroflexus aurantiacus is a photosynthetic bacterium isolated from hot springs, belonging to the green non-sulfur bacteria. This organism is thermophilic and can grow at temperatures from 35 °C to 70 °C. Chloroflexus aurantiacus can survive in the dark if oxygen is available. When grown in the dark, Chloroflexus aurantiacus has a dark orange color. When grown in sunlight it is dark green. The individual bacteria tend to form filamentous colonies enclosed in sheaths, which are known as trichomes.

C<sub>3</sub> carbon fixation Series of interconnected biochemical reactions

C3 carbon fixation is the most common of three metabolic pathways for carbon fixation in photosynthesis, the other two being C4 and CAM. This process converts carbon dioxide and ribulose bisphosphate (RuBP, a 5-carbon sugar) into two molecules of 3-phosphoglycerate through the following reaction:

<span class="mw-page-title-main">Biological carbon fixation</span> Series of interconnected biochemical reactions

Biological carbon fixation or сarbon assimilation is the process by which inorganic carbon is converted to organic compounds by living organisms. The compounds are then used to store energy and as structure for other biomolecules. Carbon is primarily fixed through photosynthesis, but some organisms use a process called chemosynthesis in the absence of sunlight.

<span class="mw-page-title-main">Calvin cycle</span> Light-independent reactions in photosynthesis

The Calvin cycle, light-independent reactions, bio synthetic phase, dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis is a series of chemical reactions that convert carbon dioxide and hydrogen-carrier compounds into glucose. The Calvin cycle is present in all photosynthetic eukaryotes and also many photosynthetic bacteria. In plants, these reactions occur in the stroma, the fluid-filled region of a chloroplast outside the thylakoid membranes. These reactions take the products of light-dependent reactions and perform further chemical processes on them. The Calvin cycle uses the chemical energy of ATP and reducing power of NADPH from the light dependent reactions to produce sugars for the plant to use. These substrates are used in a series of reduction-oxidation (redox) reactions to produce sugars in a step-wise process; there is no direct reaction that converts several molecules of CO2 to a sugar. There are three phases to the light-independent reactions, collectively called the Calvin cycle: carboxylation, reduction reactions, and ribulose 1,5-bisphosphate (RuBP) regeneration.

The light compensation point (Ic) is the light intensity on the light curve where the rate of photosynthesis exactly matches the rate of cellular respiration. At this point, the uptake of CO2 through photosynthetic pathways is equal to the respiratory release of carbon dioxide, and the uptake of O2 by respiration is equal to the photosynthetic release of oxygen. The concept of compensation points in general may be applied to other photosynthetic variables, the most important being that of CO2 concentration – CO2 compensation point (Γ).Interval of time in day time when light intensity is low due to which net gaseous exchange is zero is called as compensation point.

The photosynthetic efficiency is the fraction of light energy converted into chemical energy during photosynthesis in green plants and algae. Photosynthesis can be described by the simplified chemical reaction

Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.

<span class="mw-page-title-main">Phosphoenolpyruvate carboxylase</span> Class of enzymes

Phosphoenolpyruvate carboxylase (also known as PEP carboxylase, PEPCase, or PEPC; EC 4.1.1.31, PDB ID: 3ZGE) is an enzyme in the family of carboxy-lyases found in plants and some bacteria that catalyzes the addition of bicarbonate (HCO3−) to phosphoenolpyruvate (PEP) to form the four-carbon compound oxaloacetate and inorganic phosphate:

Malate dehydrogenase (oxaloacetate-decarboxylating) (NADP<sup>+</sup>) Enzyme

Malate dehydrogenase (oxaloacetate-decarboxylating) (NADP+) (EC 1.1.1.40) or NADP-malic enzyme (NADP-ME) is an enzyme that catalyzes the chemical reaction in the presence of a bivalent metal ion:

<span class="mw-page-title-main">Fractionation of carbon isotopes in oxygenic photosynthesis</span>

Photosynthesis converts carbon dioxide to carbohydrates via several metabolic pathways that provide energy to an organism and preferentially react with certain stable isotopes of carbon. The selective enrichment of one stable isotope over another creates distinct isotopic fractionations that can be measured and correlated among oxygenic phototrophs. The degree of carbon isotope fractionation is influenced by several factors, including the metabolism, anatomy, growth rate, and environmental conditions of the organism. Understanding these variations in carbon fractionation across species is useful for biogeochemical studies, including the reconstruction of paleoecology, plant evolution, and the characterization of food chains.

<span class="mw-page-title-main">Kinetic isotope effects of RuBisCO</span>

The kinetic isotope effect (KIE) of ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCO) is the isotopic fractionation associated solely with the step in the Calvin-Benson cycle where a molecule of carbon dioxide is attached to the 5-carbon sugar ribulose-1,5-bisphosphate (RuBP) to produce two 3-carbon sugars called 3-phosphoglycerate. This chemical reaction is catalyzed by the enzyme RuBisCO, and this enzyme-catalyzed reaction creates the primary kinetic isotope effect of photosynthesis. It is also largely responsible for the isotopic compositions of photosynthetic organisms and the heterotrophs that eat them. Understanding the intrinsic KIE of RuBisCO is of interest to earth scientists, botanists, and ecologists because this isotopic biosignature can be used to reconstruct the evolution of photosynthesis and the rise of oxygen in the geologic record, reconstruct past evolutionary relationships and environmental conditions, and infer plant relationships and productivity in modern environments.

<span class="mw-page-title-main">2-Phosphoglycolate</span> Chemical compound

2-Phosphoglycolate (chemical formula C2H2O6P3-; also known as phosphoglycolate, 2-PG, or PG) is a natural metabolic product of the oxygenase reaction mediated by the enzyme ribulose 1,5-bisphosphate carboxylase (RuBisCo).

<span class="mw-page-title-main">Photoautotrophism</span> Organisms that use light and inorganic carbon to produce organic materials

Photoautotrophs are organisms that can utilize light energy from sunlight and elements from inorganic compounds to produce organic materials needed to sustain their own metabolism. This biological activity is known as photosynthesis, and examples of such photosynthetic organisms include plants, algae and cyanobacteria.

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