89-479: See text . Physematium is a genus of ferns in the family Woodsiaceae . It was treated as a synonym of Woodsia until 2020 when a molecular phylogenetic study showed its distinctiveness. As of June 2023, Plants of the World Online continued to treat the genus as a synonym of Woodsia . As of June 2023, World Ferns accepted 20 species and two hybrids: This Polypodiales -related article
178-500: A biochemical pump that collects carbon from the organ interior (or from the soil ) and not from the atmosphere. Cyanobacteria possess carboxysomes , which increase the concentration of CO 2 around RuBisCO to increase the rate of photosynthesis. An enzyme, carbonic anhydrase , located within the carboxysome, releases CO 2 from dissolved hydrocarbonate ions (HCO 3 ). Before the CO 2 can diffuse out, RuBisCO concentrated within
267-734: A class Equisetopsida ( Embryophyta ) encompassing all land plants. This is referred to as Equisetopsida sensu lato to distinguish it from the narrower use to refer to horsetails alone, Equisetopsida sensu stricto . They placed the lycopods into subclass Lycopodiidae and the ferns, keeping the term monilophytes, into five subclasses, Equisetidae, Ophioglossidae, Psilotidae, Marattiidae and Polypodiidae, by dividing Smith's Psilotopsida into its two orders and elevating them to subclass (Ophioglossidae and Psilotidae). Christenhusz et al. (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again. Christenhusz and Chase (2014) developed
356-409: A different leaf anatomy from C 3 plants, and fix the CO 2 at night, when their stomata are open. CAM plants store the CO 2 mostly in the form of malic acid via carboxylation of phosphoenolpyruvate to oxaloacetate , which is then reduced to malate. Decarboxylation of malate during the day releases CO 2 inside the leaves, thus allowing carbon fixation to 3-phosphoglycerate by RuBisCO. CAM
445-628: A few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand ). Roots are underground non-photosynthetic structures that take up water and nutrients from soil . They are always fibrous and are structurally very similar to the roots of seed plants. As in all vascular plants , the sporophyte is the dominant phase or generation in the life cycle . The gametophytes of ferns, however, are very different from those of seed plants. They are free-living and resemble liverworts , whereas those of seed plants develop within
534-483: A new classification of ferns and lycopods. They used the term Polypodiophyta for the ferns, subdivided like Smith et al. into four groups (shown with equivalents in the Smith system), with 21 families, approximately 212 genera and 10,535 species; This was a considerable reduction in the number of families from the 37 in the system of Smith et al., since the approach was more that of lumping rather than splitting. For instance
623-930: A number of families were reduced to subfamilies. Subsequently, a consensus group was formed, the Pteridophyte Phylogeny Group (PPG), analogous to the Angiosperm Phylogeny Group , publishing their first complete classification in November 2016. They recognise ferns as a class, the Polypodiopsida, with four subclasses as described by Christenhusz and Chase, and which are phylogenetically related as in this cladogram: Equisetales Ophioglossales Psilotales Marattiales Osmundales Hymenophyllales Gleicheniales Schizaeales Photosynthesis Photosynthesis ( / ˌ f oʊ t ə ˈ s ɪ n θ ə s ɪ s / FOH -tə- SINTH -ə-sis )
712-480: A phospholipid outer membrane, and an intermembrane space. Enclosed by the membrane is an aqueous fluid called the stroma. Embedded within the stroma are stacks of thylakoids (grana), which are the site of photosynthesis. The thylakoids appear as flattened disks. The thylakoid itself is enclosed by the thylakoid membrane, and within the enclosed volume is a lumen or thylakoid space. Embedded in the thylakoid membrane are integral and peripheral membrane protein complexes of
801-479: A photocomplex. When a photon is absorbed by a chromophore, it is converted into a quasiparticle referred to as an exciton , which jumps from chromophore to chromophore towards the reaction center of the photocomplex, a collection of molecules that traps its energy in a chemical form accessible to the cell's metabolism. The exciton's wave properties enable it to cover a wider area and try out several possible paths simultaneously, allowing it to instantaneously "choose"
890-405: A photon by the antenna complex loosens an electron by a process called photoinduced charge separation . The antenna system is at the core of the chlorophyll molecule of the photosystem II reaction center. That loosened electron is taken up by the primary electron-acceptor molecule, pheophytin . As the electrons are shuttled through an electron transport chain (the so-called Z-scheme shown in
979-422: A plant's photosynthetic response. Integrated chlorophyll fluorometer – gas exchange systems allow a more precise measure of photosynthetic response and mechanisms. While standard gas exchange photosynthesis systems can measure Ci, or substomatal CO 2 levels, the addition of integrated chlorophyll fluorescence measurements allows a more precise measurement of C C, the estimation of CO 2 concentration at
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#17328447849011068-462: A protective coating called an indusium . The arrangement of the sporangia is important in classification. In monomorphic ferns, the fertile and sterile leaves look morphologically the same, and both are able to photosynthesize. In hemidimorphic ferns, just a portion of the fertile leaf is different from the sterile leaves. In dimorphic (holomorphic) ferns, the two types of leaves are morphologically distinct . The fertile leaves are much narrower than
1157-450: A redox-active tyrosine residue that is oxidized by the energy of P680 . This resets the ability of P680 to absorb another photon and release another photo-dissociated electron. The oxidation of water is catalyzed in photosystem II by a redox-active structure that contains four manganese ions and a calcium ion ; this oxygen-evolving complex binds two water molecules and contains the four oxidizing equivalents that are used to drive
1246-430: A short-lived structure anchored to the ground by rhizoids called gametophyte which produce gametes. When a mature fertile frond bears sori, and spores are released, the spores will settle on the soil and send out rhizoids , while it develops into a prothallus . The prothallus bears spherical antheridia ( s.g. antheridium ) which produce antherozoids (male gametophytes) and archegonia ( s.g. archegonium ) which release
1335-812: A significant input to the nitrogen nutrition of rice paddies . They also play certain roles in folklore. Extant ferns are herbaceous perennials and most lack woody growth. When woody growth is present, it is found in the stem. Their foliage may be deciduous or evergreen , and some are semi-evergreen depending on the climate. Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots. Ferns differ from spermatophytes in that they reproduce by spores rather than having flowers and producing seeds. However, they also differ from spore-producing bryophytes in that, like seed plants, they are polysporangiophytes , their sporophytes branching and producing many sporangia. Also unlike bryophytes, fern sporophytes are free-living and only briefly dependent on
1424-399: A simpler method that employs a pigment similar to those used for vision in animals. The bacteriorhodopsin changes its configuration in response to sunlight, acting as a proton pump. This produces a proton gradient more directly, which is then converted to chemical energy. The process does not involve carbon dioxide fixation and does not release oxygen, and seems to have evolved separately from
1513-568: A single oosphere . The antherozoid swims up the archegonium and fertilize the oosphere, resulting in a zygote, which will grow into a separate sporophyte, while the gametophyte shortly persists as a free-living plant. Carl Linnaeus (1753) originally recognized 15 genera of ferns and fern allies, classifying them in class Cryptogamia in two groups, Filices (e.g. Polypodium ) and Musci (mosses). By 1806 this had increased to 38 genera, and has progressively increased since ( see Schuettpelz et al (2018) ). Ferns were traditionally classified in
1602-481: A source of carbon atoms to carry out photosynthesis; photoheterotrophs use organic compounds, rather than carbon dioxide, as a source of carbon. In plants, algae, and cyanobacteria, photosynthesis releases oxygen. This oxygenic photosynthesis is by far the most common type of photosynthesis used by living organisms. Some shade-loving plants (sciophytes) produce such low levels of oxygen during photosynthesis that they use all of it themselves instead of releasing it to
1691-534: A subsequent sequence of light-independent reactions called the Calvin cycle . In this process, atmospheric carbon dioxide is incorporated into already existing organic compounds, such as ribulose bisphosphate (RuBP). Using the ATP and NADPH produced by the light-dependent reactions, the resulting compounds are then reduced and removed to form further carbohydrates, such as glucose . In other bacteria, different mechanisms like
1780-405: A type of non- carbon-fixing anoxygenic photosynthesis, where the simpler photopigment retinal and its microbial rhodopsin derivatives are used to absorb green light and power proton pumps to directly synthesize adenosine triphosphate (ATP), the "energy currency" of cells. Such archaeal photosynthesis might have been the earliest form of photosynthesis that evolved on Earth, as far back as
1869-591: A wide variety of colors. These pigments are embedded in plants and algae in complexes called antenna proteins. In such proteins, the pigments are arranged to work together. Such a combination of proteins is also called a light-harvesting complex . Although all cells in the green parts of a plant have chloroplasts, the majority of those are found in specially adapted structures called leaves . Certain species adapted to conditions of strong sunlight and aridity , such as many Euphorbia and cactus species, have their main photosynthetic organs in their stems. The cells in
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#17328447849011958-493: Is polyphyletic , the term fern allies should be abandoned, except in a historical context. More recent genetic studies demonstrated that the Lycopodiophyta are more distantly related to other vascular plants , having radiated evolutionarily at the base of the vascular plant clade , while both the whisk ferns and horsetails are as closely related to leptosporangiate ferns as the ophioglossoid ferns and Marattiaceae . In fact,
2047-404: Is a stub . You can help Misplaced Pages by expanding it . Fern The ferns ( Polypodiopsida or Polypodiophyta ) are a group of vascular plants (plants with xylem and phloem ) that reproduce via spores and have neither seeds nor flowers . They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients, and in having life cycles in which
2136-590: 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 metabolism . Photosynthesis usually refers to oxygenic photosynthesis , a process that produces oxygen. Photosynthetic organisms store the chemical energy so produced within intracellular organic compounds (compounds containing carbon) like sugars, glycogen , cellulose and starches . To use this stored chemical energy, an organism's cells metabolize
2225-490: Is an endothermic redox reaction. In general outline, photosynthesis is the opposite of cellular respiration : while photosynthesis is a process of reduction of carbon dioxide to carbohydrates, cellular respiration is the oxidation of carbohydrates or other nutrients to carbon dioxide. Nutrients used in cellular respiration include carbohydrates, amino acids and fatty acids. These nutrients are oxidized to produce carbon dioxide and water, and to release chemical energy to drive
2314-498: Is both an evolutionary precursor to C 4 and a useful carbon-concentrating mechanism in its own right. Xerophytes , such as cacti and most succulents , also use PEP carboxylase to capture carbon dioxide in a process called Crassulacean acid metabolism (CAM). In contrast to C 4 metabolism, which spatially separates the CO 2 fixation to PEP from the Calvin cycle, CAM temporally separates these two processes. CAM plants have
2403-474: Is commonly measured in μmols /( m / s ), parts per million, or volume per million; and H 2 O is commonly measured in mmols /(m /s) or in mbars . By measuring CO 2 assimilation , ΔH 2 O, leaf temperature, barometric pressure , leaf area, and photosynthetically active radiation (PAR), it becomes possible to estimate, "A" or carbon assimilation, "E" or transpiration , "gs" or stomatal conductance , and "Ci" or intracellular CO 2 . However, it
2492-430: Is converted to CO 2 by an oxalate oxidase enzyme, and the produced CO 2 can support the Calvin cycle reactions. Reactive hydrogen peroxide (H 2 O 2 ), the byproduct of oxalate oxidase reaction, can be neutralized by catalase . Alarm photosynthesis represents a photosynthetic variant to be added to the well-known C4 and CAM pathways. However, alarm photosynthesis, in contrast to these pathways, operates as
2581-419: Is freed from its locked position through a classic "hop". The movement of the electron towards the photo center is therefore covered in a series of conventional hops and quantum walks. Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old. More recent studies also suggest that photosynthesis may have begun about 3.4 billion years ago, though
2670-412: Is further excited by the light absorbed by that photosystem . The electron is then passed along a chain of electron acceptors to which it transfers some of its energy . The energy delivered to the electron acceptors is used to move hydrogen ions across the thylakoid membrane into the lumen . The electron is eventually used to reduce the coenzyme NADP with an H to NADPH (which has functions in
2759-578: Is intermediate between the eusporangiate ferns and the leptosporangiate ferns. Rai and Graham (2010) broadly supported the primary groups, but queried their relationships, concluding that "at present perhaps the best that can be said about all relationships among the major lineages of monilophytes in current studies is that we do not understand them very well". Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato , but also suggested that uncertainties remained in their precise placement. Other classifications have raised Ophioglossales to
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2848-451: Is more common to use chlorophyll fluorescence for plant stress measurement , where appropriate, because the most commonly used parameters FV/FM and Y(II) or F/FM' can be measured in a few seconds, allowing the investigation of larger plant populations. Gas exchange systems that offer control of CO 2 levels, above and below ambient , allow the common practice of measurement of A/Ci curves, at different CO 2 levels, to characterize
2937-406: Is passed through a food chain . The fixation or reduction of carbon dioxide is a process in which carbon dioxide combines with a five-carbon sugar , ribulose 1,5-bisphosphate , to yield two molecules of a three-carbon compound, glycerate 3-phosphate , also known as 3-phosphoglycerate. Glycerate 3-phosphate, in the presence of ATP and NADPH produced during the light-dependent stages,
3026-623: Is reduced to glyceraldehyde 3-phosphate . This product is also referred to as 3-phosphoglyceraldehyde (PGAL) or, more generically, as triose phosphate. Most (five out of six molecules) of the glyceraldehyde 3-phosphate produced are used to regenerate ribulose 1,5-bisphosphate so the process can continue. The triose phosphates not thus "recycled" often condense to form hexose phosphates, which ultimately yield sucrose , starch , and cellulose , as well as glucose and fructose . The sugars produced during carbon metabolism yield carbon skeletons that can be used for other metabolic reactions like
3115-429: Is then translocated to specialized bundle sheath cells where the enzyme RuBisCO and other Calvin cycle enzymes are located, and where CO 2 released by decarboxylation of the four-carbon acids is then fixed by RuBisCO activity to the three-carbon 3-phosphoglyceric acids . The physical separation of RuBisCO from the oxygen-generating light reactions reduces photorespiration and increases CO 2 fixation and, thus,
3204-404: Is then converted into the final carbohydrate products. The simple carbon sugars photosynthesis produces are then used to form other organic compounds , such as the building material cellulose , the precursors for lipid and amino acid biosynthesis, or as a fuel in cellular respiration . The latter occurs not only in plants but also in animals when the carbon and energy from plants
3293-499: Is used by 16,000 species of plants. Calcium-oxalate -accumulating plants, such as Amaranthus hybridus and Colobanthus quitensis , show a variation of photosynthesis where calcium oxalate crystals function as dynamic carbon pools , supplying carbon dioxide (CO 2 ) to photosynthetic cells when stomata are partially or totally closed. This process was named alarm photosynthesis . Under stress conditions (e.g., water deficit ), oxalate released from calcium oxalate crystals
3382-474: Is vital for climate processes, as it captures carbon dioxide from the air and binds it into plants, harvested produce and soil. Cereals alone are estimated to bind 3,825 Tg or 3.825 Pg of carbon dioxide every year, i.e. 3.825 billion metric tons. Most photosynthetic organisms are photoautotrophs , which means that they are able to synthesize food directly from carbon dioxide and water using energy from light. However, not all organisms use carbon dioxide as
3471-640: The Cretaceous , contemporaneous with the rise of flowering plants that came to dominate the world's flora. Ferns are not of major economic importance, but some are used for food, medicine, as biofertilizer , as ornamental plants, and for remediating contaminated soil. They have been the subject of research for their ability to remove some chemical pollutants from the atmosphere. Some fern species, such as bracken ( Pteridium aquilinum ) and water fern ( Azolla filiculoides ), are significant weeds worldwide. Some fern genera, such as Azolla , can fix nitrogen and make
3560-572: The Paleoarchean , preceding that of cyanobacteria (see Purple Earth hypothesis ). While the details may differ between species , the process always begins when light energy is absorbed by the reaction centers , proteins that contain photosynthetic pigments or chromophores . In plants, these proteins are chlorophylls (a porphyrin derivative that absorbs the red and blue spectrums of light, thus reflecting green) held inside chloroplasts , abundant in leaf cells. In bacteria, they are embedded in
3649-472: The class Filices, and later in a Division of the Plant Kingdom named Pteridophyta or Filicophyta. Pteridophyta is no longer recognised as a valid taxon because it is paraphyletic . The ferns are also referred to as Polypodiophyta or, when treated as a subdivision of Tracheophyta (vascular plants), Polypodiopsida, although this name sometimes only refers to leptosporangiate ferns. Traditionally, all of
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3738-637: The light reaction of photosynthesis by using chlorophyll fluorometers . Actual plants' photosynthetic efficiency varies with the frequency of the light being converted, light intensity , temperature , and proportion of carbon dioxide in the atmosphere , and can vary from 0.1% to 8%. By comparison, solar panels convert light into electric energy at an efficiency of approximately 6–20% for mass-produced panels, and above 40% in laboratory devices. Scientists are studying photosynthesis in hopes of developing plants with increased yield . The efficiency of both light and dark reactions can be measured, but
3827-434: The light-independent (or "dark") reactions, the enzyme RuBisCO captures CO 2 from the atmosphere and, in a process called the Calvin cycle , uses the newly formed NADPH and releases three-carbon sugars , which are later combined to form sucrose and starch . The overall equation for the light-independent reactions in green plants is Carbon fixation produces the three-carbon sugar intermediate , which
3916-417: The palisade mesophyll cells where most of the photosynthesis takes place. In the light-dependent reactions , one molecule of the pigment chlorophyll absorbs one photon and loses one electron . This electron is taken up by a modified form of chlorophyll called pheophytin , which passes the electron to a quinone molecule, starting the flow of electrons down an electron transport chain that leads to
4005-417: The photosynthetic capacity of the leaf . C 4 plants can produce more sugar than C 3 plants in conditions of high light and temperature . Many important crop plants are C 4 plants, including maize , sorghum , sugarcane , and millet . Plants that do not use PEP-carboxylase in carbon fixation are called C 3 plants because the primary carboxylation reaction , catalyzed by RuBisCO, produces
4094-462: The photosystems , quantum efficiency and the CO 2 assimilation rates. With some instruments, even wavelength dependency of the photosynthetic efficiency can be analyzed . A phenomenon known as quantum walk increases the efficiency of the energy transport of light significantly. In the photosynthetic cell of an alga , bacterium , or plant, there are light-sensitive molecules called chromophores arranged in an antenna-shaped structure called
4183-437: The plasma membrane . In these light-dependent reactions, some energy is used to strip electrons from suitable substances, such as water, producing oxygen gas. The hydrogen freed by the splitting of water is used in the creation of two important molecules that participate in energetic processes: reduced nicotinamide adenine dinucleotide phosphate (NADPH) and ATP. In plants, algae, and cyanobacteria, sugars are synthesized by
4272-450: The reverse Krebs cycle are used to achieve the same end. The first photosynthetic organisms probably evolved early in the evolutionary history of life using reducing agents such as hydrogen or hydrogen sulfide, rather than water, as sources of electrons. Cyanobacteria appeared later; the excess oxygen they produced contributed directly to the oxygenation of the Earth , which rendered
4361-433: The CO 2 concentration in the leaves under these conditions. Plants that use the C 4 carbon fixation process chemically fix carbon dioxide in the cells of the mesophyll by adding it to the three-carbon molecule phosphoenolpyruvate (PEP), a reaction catalyzed by an enzyme called PEP carboxylase , creating the four-carbon organic acid oxaloacetic acid . Oxaloacetic acid or malate synthesized by this process
4450-419: The action spectrum is blue-green light, which allows these algae to use the blue end of the spectrum to grow in the deeper waters that filter out the longer wavelengths (red light) used by above-ground green plants. The non-absorbed part of the light spectrum is what gives photosynthetic organisms their color (e.g., green plants, red algae, purple bacteria ) and is the least effective for photosynthesis in
4539-521: The atmosphere. Although there are some differences between oxygenic photosynthesis in plants , algae , and cyanobacteria , the overall process is quite similar in these organisms. There are also many varieties of anoxygenic photosynthesis , used mostly by bacteria, which consume carbon dioxide but do not release oxygen. Carbon dioxide is converted into sugars in a process called carbon fixation ; photosynthesis captures energy from sunlight to convert carbon dioxide into carbohydrates . Carbon fixation
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#17328447849014628-478: The branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls that are more complex than the microphylls of clubmosses . Most ferns are leptosporangiate ferns . They produce coiled fiddleheads that uncoil and expand into fronds . The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida , comprising both
4717-480: The carboxysome quickly sponges it up. HCO 3 ions are made from CO 2 outside the cell by another carbonic anhydrase and are actively pumped into the cell by a membrane protein. They cannot cross the membrane as they are charged, and within the cytosol they turn back into CO 2 very slowly without the help of carbonic anhydrase. This causes the HCO 3 ions to accumulate within the cell from where they diffuse into
4806-491: The carboxysomes. Pyrenoids in algae and hornworts also act to concentrate CO 2 around RuBisCO. The overall process of photosynthesis takes place in four stages: Plants usually convert light into chemical energy with a photosynthetic efficiency of 3–6%. Absorbed light that is unconverted is dissipated primarily as heat , with a small fraction (1–2%) reemitted as chlorophyll fluorescence at longer (redder) wavelengths . This fact allows measurement of
4895-409: The conditions of non-cyclic electron flow in green plants is: Not all wavelengths of light can support photosynthesis. The photosynthetic action spectrum depends on the type of accessory pigments present. For example, in green plants , the action spectrum resembles the absorption spectrum for chlorophylls and carotenoids with absorption peaks in violet-blue and red light. In red algae ,
4984-503: The diagram), a chemiosmotic potential is generated by pumping proton cations ( H ) across the membrane and into the thylakoid space . An ATP synthase enzyme uses that chemiosmotic potential to make ATP during photophosphorylation , whereas NADPH is a product of the terminal redox reaction in the Z-scheme . The electron enters a chlorophyll molecule in Photosystem I . There it
5073-505: The equation for this process is: This equation emphasizes that water is both a reactant in the light-dependent reaction and a product of the light-independent reaction , but canceling n water molecules from each side gives the net equation: Other processes substitute other compounds (such as arsenite ) for water in the electron-supply role; for example some microbes use sunlight to oxidize arsenite to arsenate : The equation for this reaction is: Photosynthesis occurs in two stages. In
5162-518: The evolution of complex life possible. The average rate of energy captured by global photosynthesis is approximately 130 terawatts , which is about eight times the total power consumption of human civilization . Photosynthetic organisms also convert around 100–115 billion tons (91–104 Pg petagrams , or billions of metric tons), of carbon into biomass per year. Photosynthesis was discovered in 1779 by Jan Ingenhousz . He showed that plants need light, not just air, soil, and water. Photosynthesis
5251-582: The first stage, light-dependent reactions or light reactions capture the energy of light and use it to make the hydrogen carrier NADPH and the energy-storage molecule ATP . During the second stage, the light-independent reactions use these products to capture and reduce carbon dioxide. Most organisms that use oxygenic photosynthesis use visible light for the light-dependent reactions, although at least three use shortwave infrared or, more specifically, far-red radiation. Some organisms employ even more radical variants of photosynthesis. Some archaea use
5340-441: The first step of the Z-scheme , requires an external source of electrons to reduce its oxidized chlorophyll a reaction center. The source of electrons for photosynthesis in green plants and cyanobacteria is water. Two water molecules are oxidized by the energy of four successive charge-separation reactions of photosystem II to yield a molecule of diatomic oxygen and four hydrogen ions. The electrons yielded are transferred to
5429-431: The fronds are branched more than once, it can also be a combination of the pinnatifid are pinnate shapes. If the leaf blades are divided twice, the plant has bipinnate fronds, and tripinnate fronds if they branch three times, and all the way to tetra- and pentapinnate fronds. In tree ferns, the main stalk that connects the leaf to the stem (known as the stipe), often has multiple leaflets. The leafy structures that grow from
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#17328447849015518-488: The inclusion of Equisetaceae in the ferns, notably relating to the construction of their sperm and peculiarities of their roots. The leptosporangiate ferns are sometimes called "true ferns". This group includes most plants familiarly known as ferns. Modern research supports older ideas based on morphology that the Osmundaceae diverged early in the evolutionary history of the leptosporangiate ferns; in certain ways this family
5607-399: The interior of a cell, giving the membrane a very large surface area and therefore increasing the amount of light that the bacteria can absorb. In plants and algae, photosynthesis takes place in organelles called chloroplasts . A typical plant cell contains about 10 to 100 chloroplasts. The chloroplast is enclosed by a membrane. This membrane is composed of a phospholipid inner membrane,
5696-433: The interior tissues of a leaf, called the mesophyll , can contain between 450,000 and 800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is coated with a water-resistant waxy cuticle that protects the leaf from excessive evaporation of water and decreases the absorption of ultraviolet or blue light to minimize heating . The transparent epidermis layer allows light to pass through to
5785-446: The leptosporangiate ( Polypodiidae ) and eusporangiate ferns , the latter group including horsetails , whisk ferns , marattioid ferns , and ophioglossoid ferns . The fern crown group , consisting of the leptosporangiates and eusporangiates, is estimated to have originated in the late Silurian period 423.2 million years ago, but Polypodiales , the group that makes up 80% of living fern diversity, did not appear and diversify until
5874-467: The leptosporangiate ferns. The Marattiaceae are a primitive group of tropical ferns with large, fleshy rhizomes and are now thought to be a sibling taxon to the leptosporangiate ferns. Several other groups of species were considered fern allies: the clubmosses , spikemosses , and quillworts in Lycopodiophyta ; the whisk ferns of Psilotaceae ; and the horsetails of Equisetaceae . Since this grouping
5963-434: The light reaction, and infrared gas analyzers can measure the dark reaction . An integrated chlorophyll fluorometer and gas exchange system can investigate both light and dark reactions when researchers use the two separate systems together. Infrared gas analyzers and some moisture sensors are sensitive enough to measure the photosynthetic assimilation of CO 2 and of Δ H 2 O using reliable methods . CO 2
6052-433: The light-independent reaction); at that point, the path of that electron ends. The cyclic reaction is similar to that of the non-cyclic but differs in that it generates only ATP, and no reduced NADP (NADPH) is created. The cyclic reaction takes place only at photosystem I. Once the electron is displaced from the photosystem, the electron is passed down the electron acceptor molecules and returns to photosystem I, from where it
6141-555: The maternal gametophyte . The green , photosynthetic part of the plant is technically a megaphyll and in ferns, it is often called a frond . New leaves typically expand by the unrolling of a tight spiral called a crozier or fiddlehead into fronds . This uncurling of the leaf is termed circinate vernation . Leaves are divided into two types: sporophylls and tropophylls. Sporophylls produce spores; tropophylls do not. Fern spores are borne in sporangia which are usually clustered to form sori . The sporangia may be covered with
6230-432: The more common types of photosynthesis. In photosynthetic bacteria, the proteins that gather light for photosynthesis are embedded in cell membranes . In its simplest form, this involves the membrane surrounding the cell itself. However, the membrane may be tightly folded into cylindrical sheets called thylakoids , or bunched up into round vesicles called intracytoplasmic membranes . These structures can fill most of
6319-439: The most efficient route, where it will have the highest probability of arriving at its destination in the minimum possible time. Because that quantum walking takes place at temperatures far higher than quantum phenomena usually occur, it is only possible over very short distances. Obstacles in the form of destructive interference cause the particle to lose its wave properties for an instant before it regains them once again after it
6408-498: The organic compounds through cellular respiration . Photosynthesis plays a critical role in 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. Some bacteria also perform anoxygenic photosynthesis , which uses bacteriochlorophyll to split hydrogen sulfide as a reductant instead of water, producing sulfur instead of oxygen. Archaea such as Halobacterium also perform
6497-410: The organism's metabolism . Photosynthesis and cellular respiration are distinct processes, as they take place through different sequences of chemical reactions and in different cellular compartments (cellular respiration in mitochondria ). The general equation for photosynthesis as first proposed by Cornelis van Niel is: Since water is used as the electron donor in oxygenic photosynthesis,
6586-556: The photosynthetic system. Plants absorb light primarily using the pigment chlorophyll . The green part of the light spectrum is not absorbed but is reflected, which is the reason that most plants have a green color. Besides chlorophyll, plants also use pigments such as carotenes and xanthophylls . Algae also use chlorophyll, but various other pigments are present, such as phycocyanin , carotenes , and xanthophylls in green algae , phycoerythrin in red algae (rhodophytes) and fucoxanthin in brown algae and diatoms resulting in
6675-476: The production of amino acids and lipids . In hot and dry conditions , plants close their stomata to prevent water loss. Under these conditions, CO 2 will decrease and oxygen gas , produced by the light reactions of photosynthesis, will increase, causing an increase of photorespiration by the oxygenase activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and decrease in carbon fixation. Some plants have evolved mechanisms to increase
6764-430: The rank of a fifth class, separating the whisk ferns and ophioglossoid ferns. The ferns are related to other groups as shown in the following cladogram: Lycophytes [REDACTED] Ferns [REDACTED] Gymnosperms [REDACTED] Angiosperms [REDACTED] The classification of Smith et al. in 2006 treated ferns as four classes: In addition they defined 11 orders and 37 families. That system
6853-486: The relationship between the two can be complex. For example, the light reaction creates ATP and NADPH energy molecules , which C 3 plants can use for carbon fixation or photorespiration . Electrons may also flow to other electron sinks. For this reason, it is not uncommon for authors to differentiate between work done under non-photorespiratory conditions and under photorespiratory conditions . Chlorophyll fluorescence of photosystem II can measure
6942-462: The respective organisms . In plants , light-dependent reactions occur in the thylakoid membranes of the chloroplasts where they drive the synthesis of ATP and NADPH . The light-dependent reactions are of two forms: cyclic and non-cyclic . In the non-cyclic reaction, the photons are captured in the light-harvesting antenna complexes of photosystem II by chlorophyll and other accessory pigments (see diagram at right). The absorption of
7031-479: The site of carboxylation in the chloroplast, to replace Ci. CO 2 concentration in the chloroplast becomes possible to estimate with the measurement of mesophyll conductance or g m using an integrated system. Photosynthesis measurement systems are not designed to directly measure the amount of light the leaf absorbs, but analysis of chlorophyll fluorescence , P700 - and P515-absorbance, and gas exchange measurements reveal detailed information about, e.g.,
7120-476: The spore producing vascular plants were informally denominated the pteridophytes , rendering the term synonymous with ferns and fern allies . This can be confusing because members of the division Pteridophyta were also denominated pteridophytes ( sensu stricto ). Traditionally, three discrete groups have been denominated ferns: two groups of eusporangiate ferns, the families Ophioglossaceae ( adder's tongues , moonworts , and grape ferns) and Marattiaceae ; and
7209-417: The spore wall and are dependent on the parent sporophyte for their nutrition. A fern gametophyte typically consists of: The lifecycle of a fern involves two stages, as in club mosses and horsetails . In stage one, the spores are produced by sporophytes in sporangia , which are clustered together in sori ( s.g. sorus ), developing on the underside of fertile fronds. In stage two, the spores germinate into
7298-490: The sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae . The anatomy of fern leaves can be anywhere from simple to highly divided, or even indeterminate (e.g. Gleicheniaceae , Lygodiaceae ). The divided forms are pinnate , where the leaf segments are completely separated from one other, or pinnatifid (partially pinnate), where the leaf segments are still partially connected. When
7387-463: The stipe are known as pinnae and are often again divided into smaller pinnules. Fern stems are often loosely called rhizomes , even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping stolons (e.g., Polypodiaceae ), and many groups have above-ground erect semi-woody trunks (e.g., Cyatheaceae , the scaly tree ferns). These can reach up to 20 meters (66 ft) tall in
7476-461: The three-carbon 3-phosphoglyceric acids directly in the Calvin-Benson cycle . Over 90% of plants use C 3 carbon fixation, compared to 3% that use C 4 carbon fixation; however, the evolution of C 4 in over sixty plant lineages makes it a striking example of convergent evolution . C 2 photosynthesis , which involves carbon-concentration by selective breakdown of photorespiratory glycine,
7565-424: The ultimate reduction of NADP to NADPH . In addition, this creates a proton gradient (energy gradient) across the chloroplast membrane , which is used by ATP synthase in the synthesis of ATP . The chlorophyll molecule ultimately regains the electron it lost when a water molecule is split in a process called photolysis , which releases oxygen . The overall equation for the light-dependent reactions under
7654-408: The water-oxidizing reaction (Kok's S-state diagrams). The hydrogen ions are released in the thylakoid lumen and therefore contribute to the transmembrane chemiosmotic potential that leads to ATP synthesis . Oxygen is a waste product of light-dependent reactions, but the majority of organisms on Earth use oxygen and its energy for cellular respiration , including photosynthetic organisms . In
7743-477: The whisk ferns and ophioglossoid ferns are demonstrably a clade , and the horsetails and Marattiaceae are arguably another clade. Smith et al. (2006) carried out the first higher-level pteridophyte classification published in the molecular phylogenetic era, and considered the ferns as monilophytes, as follows: Molecular data, which remain poorly constrained for many parts of the plants' phylogeny, have been supplemented by morphological observations supporting
7832-844: Was a consensus of a number of studies, and was further refined. The phylogenetic relationships are shown in the following cladogram (to the level of orders). This division into four major clades was then confirmed using morphology alone. Lycopodiophytes (club mosses, spike mosses, quillworts) Spermatophytes (seed plants) Equisetales (horsetails) [REDACTED] Ophioglossales (grapeferns etc.) Psilotales (whisk ferns) [REDACTED] Marattiales [REDACTED] Osmundales [REDACTED] Hymenophyllales (filmy ferns) [REDACTED] Gleicheniales [REDACTED] Schizaeales Salviniales (heterosporous) Cyatheales (tree ferns) [REDACTED] Polypodiales [REDACTED] Subsequently, Chase and Reveal considered both lycopods and ferns as subclasses of
7921-462: Was emitted, hence the name cyclic reaction . Linear electron transport through a photosystem will leave the reaction center of that photosystem oxidized . Elevating another electron will first require re-reduction of the reaction center. The excited electrons lost from the reaction center ( P700 ) of photosystem I are replaced by transfer from plastocyanin , whose electrons come from electron transport through photosystem II . Photosystem II, as
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