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Carbon fixation
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For other uses, see Carbon cycle and Carbon sequestration.
Cyanobacteria such as these carry out photosynthesis. Their emergence foreshadowed the evolution of many photosynthetic plants, which oxygenated Earth's atmosphere.
Carbon fixation or сarbon assimilation is the conversion process of inorganic carbon (carbon dioxide) to organic compounds by living organisms. The most prominent example is photosynthesis, although chemosynthesis is another form of carbon fixation that can take place in the absence of sunlight. Organisms that grow by fixing carbon are called autotrophs. Autotrophs include photoautotrophs, which synthesize organic compounds using the energy of sunlight, and lithoautotrophs, which synthesize organic compounds using the energy of inorganic oxidation. Heterotrophs are organisms that grow using the carbon fixed by autotrophs. The organic compounds are used by heterotrophs to produce energy and to build body structures. "Fixed carbon", "reduced carbon", and "organic carbon" are equivalent terms for various organic compounds.[1]
Contents
[hide]
1 Net vs gross CO2 fixation
2 Overview of pathways
3 Oxygenic photosynthesis
3.1 Evolutionary considerations
3.2 Carbon concentrating mechanisms
3.2.1 CAM plants
3.2.2 C4 plants
3.2.3 C3 plants
4 Other autotrophic pathways
4.1 Reductive citric acid cycle
4.2 Reductive acetyl CoA pathway
4.3 3-Hydroxypropionate and two related cycles
5 Chemosynthesis
6 Non-autotrophic pathways
7 Carbon isotope discrimination
8 References
9 Further reading
9.1 Descent of plants and algae
Net vs gross CO2 fixation[edit]
Graphic showing net annual amounts of CO2 fixation by land and sea-based organisms.
It is estimated that approximately 258 billion tons of carbon dioxide are converted by photosynthesis annually. The majority of the fixation occurs in marine environments, especially areas of high nutrients. The gross amount of carbon dioxide fixed is much larger since approximately 40% is consumed by respiration following photosynthesis.[1] Given the scale of this process, it is understandable that RuBisCO is the most abundant protein on Earth.
Overview of pathways[edit]
Six autotrophic carbon fixation pathways are known as of 2011. The Calvin cycle fixes carbon in the chloroplasts of plants and algae, and in the cyanobacteria. It also fixes carbon in the anoxygenic photosynthetic proteobacteria called purple bacteria, and in some non-phototrophic proteobacteria.[2]
Oxygenic photosynthesis[edit]
In photosynthesis, energy from sunlight drives the carbon fixation pathway. Oxygenic photosynthesis is used by the primary producers—plants, algae, and cyanobacteria. They contain the pigment chlorophyll, and use the Calvin cycle to fix carbon autotrophically. The process works like this:
2H2O → 4e− + 4H+ + O2
CO2 + 4e− + 4H+ → CH2O + H2O
In the first step, water is dissociated into electrons, protons, and free oxygen. This allows the use of water, one of the most abundant substances on Earth, as an electron donor—as a source of reducing power. The release of free oxygen is a side-effect of enormous consequence. The first step uses the energy of sunlight to oxidize water to O2, and, ultimately, to produce ATP
ADP + Pi ⇌ ATP + H2O
and the reductant, NADPH
NADP+ + 2e− + 2H+ ⇌ NADPH + H+
In the second step, called the Calvin cycle, the actual fixation of carbon dioxide is carried out. This process consumes ATP and NADPH. The Calvin cycle in plants accounts for the preponderance of carbon fixation on land. In algae and cyanobacteria, it accounts for the preponderance of carbon fixation in the oceans. The Calvin cycle converts carbon dioxide into sugar, as triose phosphate (TP), which is glyceraldehyde 3-phosphate (GAP) together with dihydroxyacetone phosphate (DHAP):
3 CO2 + 12 e− + 12 H+ + Pi → TP + 4 H2O
An alternative perspective accounts for NADPH (source of e−) and ATP:
3 CO2 + 6 NADPH + 6 H+ + 9 ATP + 5 H2O → TP + 6 NADP+ + 9 ADP + 8 Pi
The formula for inorganic phosphate (Pi) is HOPO32− + 2H+. Formulas for triose and TP are C2H3O2-CH2OH and C2H3O2-CH2OPO32− + 2H+
Evolutionary considerations[edit]
Somewhere between 3.5 and 2.3 billion years ago, the ancestors of cyanobacteria evolved oxygenic photosynthesis, enabling the use of the abundant yet relatively oxidized molecule H2O as an electron donor to the electron transport chain of light-catalyzed proton-pumping responsible for efficient ATP synthesis.[3][4] When this evolutionary breakthrough occurred, autotrophy (growth using inorganic carbon as the sole carbon source) is believed to have already been developed. However, the proliferation of cyanobacteria, due to their novel ability to exploit water as a source of electrons, radically altered the global environment by oxygenating the atmosphere and by achieving large fluxes of CO2 consumption.[5]
Carbon concentrating mechanisms[edit]
Many photosynthetic organisms have not acquired inorganic carbon concentrating mechanisms (CCM), which increase the concentration of carbon dioxide available to the initial carboxylase of the Calvin cycle, the enzyme RuBisCO. The benefits of CCM include increased tolerance to low external concentrations of inorganic carbon, and reduced losses to photorespiration. CCM can make plants more tolerant of heat and water stress.
Carbon concentrating mechanisms use the enzyme carbonic anhydrase (CA), which catalyze both the dehydration of bicarbonate to carbon dioxide and the hydration of carbon dioxide to bicarbonate
HCO3− + H+ ⇌ CO2 + H2O
Lipid membranes are much less permeable to bicarbonate than to carbon dioxide. To capture inorganic carbon more effectively, some plants have adapted the anaplerotic reactions
HCO3− + H+ + PEP → OAA + Pi
catalyzed by PEP carboxylase (PEPC), to carboxylate phosphoenolpyruvate (PEP) to oxaloacetate (OAA) which is a C4 dicarboxylic acid.
CAM plants[edit]
CAM plants that use Crassulacean acid metabolism as an adaptation for arid conditions. CO2 enters through the stomata during the night and is converted into the 4-carbon compound, malic acid, which releases CO2 for use in the Calvin cycle during the day, when the stomata are closed. The dung jade plant (Crassula ovata) and cacti are typical of CAM plants. Sixteen thousand species of plants use CAM.[6] These plants have a carbon isotope signature of −20 to −10 ‰.[7]
C4 plants[edit]
C4 plants preface the Calvin cycle with reactions that incorporate CO2 into one of the 4-carbon compounds, malic acid or aspartic acid. C4 plants have a distinctive internal leaf anatomy. Tropical grasses, such as sugar cane and maize are C4 plants, but there are many broadleaf plants that are C4. Overall, 7600 species of terrestrial plants use C4 carbon fixation, representing around 3% of all species.[8] These plants have a carbon isotope signature of −16 to −10 ‰.[7]
C3 plants[edit]
The large majority of plants are C3 plants. They are so-called to distinguish them from the CAM and C4 plants, and because the carboxylation products of the Calvin cycle are 3-carbon compounds. They lack C4 dicarboxylic acid cycles, and therefore have higher carbon dioxide compensation points than CAM or C4 plants. C3 plants have a carbon isotope signature of −24 to −33‰.[7]
Other autotrophic pathways[edit]
Of the five other autotrophic pathways, two are known only in bacteria, two only in archaea, and one in both bacteria and archaea.
Reductive citric acid cycle[edit]
The reductive citric acid cycle is the oxidative citric acid cycle run in reverse. It has been found in anaerobic and microaerobic bacteria. It was proposed in 1966 by Evans, Buchanan and Arnon who were working with the anoxygenic photosynthetic green sulfur bacterium that they called Chlorobium thiosulfatophilum. The reductive citric acid cycle is sometimes called the Arnon-Buchanan cycle.[9]
Reductive acetyl CoA pathway[edit]
The reductive acetyl CoA pathway operated in strictly anaerobic bacteria (acetogens) and archaea (methanogens). The pathway was proposed in 1965 by Ljungdahl and Wood. They were working with the gram-positive acetic acid producing bacterium Clostridium thermoaceticum, which is now named Moorella thermoacetica. Hydrogenotrophic methanogenesis, which is only found in certain archaea and accounts for 80% of global methanogenesis, is also based on the reductive acetyl CoA pathway. The pathway is often referred to as the Wood–Ljungdahl pathway.[10][11]
3-Hydroxypropionate and two related cycles[edit]
The 3-hydroxypropionate cycle is utilized only by green nonsulfur bacteria. It was proposed in 2002 for the anoxygenic photosynthetic Chloroflexus aurantiacus. None of the enzymes that participate in the 3-hydroxypropionate cycle are especially oxygen sensitive.[12][13]
A variant of the 3-hydroxypropionate pathway was found to operate in aerobic extreme thermoacidophile archaeon Metallosphaera sedula. This pathway, called the 3-hydroxypropionate/4-hydroxybutyrate cycle .[14] And yet another variant of the 3-hydroxypropionate pathway is the dicarboxylate/4-hydroxybutyrate cycle. It was discovered in anaerobic archaea. It was proposed in 2008 for the hyperthermophile archeon Ignicoccus hospitalis.[15]
Chemosynthesis[edit]
Chemosynthesis is carbon fixation driven by the oxidation of inorganic substances (e.g., hydrogen gas or hydrogen sulfide). Sulfur- and hydrogen-oxidizing bacteria often use the Calvin cycle or the reductive citric acid cycle.[16]
Non-autotrophic pathways[edit]
Although almost all heterotrophs cannot synthesize complete organic molecules from carbon dioxide, some carbon dioxide is incorporated in their metabolism.[17] Notably pyruvate carboxylase consumes carbon dioxide (as bicarbonate ions) as part of gluconeogenesis, and carbon dioxide is consumed in various anaplerotic reactions.
Carbon isotope discrimination[edit]
Some carboxylases, particularly RuBisCO, preferentially bind the lighter carbon stable isotope carbon-12 over the heavier carbon-13. This is known as carbon isotope discrimination and results in carbon-12 to carbon-13 ratios in the plant that are higher than in the free air. Measurement of this ratio is important in the evaluation of water use efficiency in plants, and also in assessing the possible or likely sources of carbon in global carbon cycle studies.
References[edit]
^ Jump up to: a b Geider, R. J., et al., "Primary productivity of planet earth: biological determinants and physical constraints in terrestrial and aquatic habitats", Global Change Biol. 2001, 7, 849–882. doi:10.1046/j.1365-2486.2001.00448.x
Jump up ^ Swan BK, Martinez-Garcia M, Preston CM, Sczyrba A, Woyke T, Lamy D, Reinthaler T, Poulton NJ, Masland ED, Gomez ML, Sieracki ME, DeLong EF, Herndl GJ, Stepanauskas R (2011). "Potential for chemolithoautotrophy among ubiquitous bacteria lineages in the dark ocean". Science. 333 (6047): 1296–300. Bibcode:2011Sci...333.1296S. PMID 21885783. doi:10.1126/science.1203690.
Jump up ^ Brasier M, McLoughlin N, Green O, Wacey D (2006). "A fresh look at the fossil evidence for early Archaean cellular life". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 361 (1470): 887–902. PMC 1578727  . PMID 16754605. doi:10.1098/rstb.2006.1835.
Jump up ^ Tomitani A, Knoll AH, Cavanaugh CM, Ohno T (2006). "The evolutionary diversification of cyanobacteria: molecular-phylogenetic and paleontological perspectives". Proc. Natl. Acad. Sci. U.S.A. 103 (14): 5442–7. Bibcode:2006PNAS..103.5442T. PMC 1459374  . PMID 16569695. doi:10.1073/pnas.0600999103.
Jump up ^ Kopp RE, Kirschvink JL, Hilburn IA, Nash CZ (2005). "The Paleoproterozoic snowball Earth: a climate disaster triggered by the evolution of oxygenic photosynthesis". Proc. Natl. Acad. Sci. U.S.A. 102 (32): 11131–6. Bibcode:2005PNAS..10211131K. PMC 1183582  . PMID 16061801. doi:10.1073/pnas.0504878102.
Jump up ^ Dodd AN, Borland AM, Haslam RP, Griffiths H, Maxwell K (2002). "Crassulacean acid metabolism: plastic, fantastic". J. Exp. Bot. 53 (369): 569–580. PMID 11886877. doi:10.1093/jexbot/53.369.569.
^ Jump up to: a b c O'Leary MH (1988). "Carbon isotopes in photosynthesis". BioScience. 38 (5): 328–336. JSTOR 1310735. doi:10.2307/1310735.
Jump up ^ Sage RF, Meirong L, Monson RK (1999). "16. The Taxonomic Distribution of C4 Photosynthesis". In Sage RF, Monson RK. C4 Plant Biology. pp. 551–580. ISBN 0-12-614440-0.
Jump up ^ Evans MC, Buchanan BB, Arnon DI (1966). "A new ferredoxin-dependent carbon reduction cycle in a photosynthetic bacterium". Proc. Natl. Acad. Sci. U.S.A. 55 (4): 928–34. Bibcode:1966PNAS...55..928E. PMC 224252  . PMID 5219700. doi:10.1073/pnas.55.4.928.
Jump up ^ Ljungdahl L, Wood HG (1965). "Incorporation of C-14 from carbon dioxide into sugar phosphates, carboxylic acids, and amino acids by Clostridium thermoaceticum". J. Bacteriol. 89: 1055–64. PMC 277595  . PMID 14276095.
Jump up ^ Ljungdahl LG (2009). "A life with acetogens, thermophiles, and cellulolytic anaerobes". Annu. Rev. Microbiol. 63: 1–25. PMID 19575555. doi:10.1146/annurev.micro.091208.073617.
Jump up ^ Herter S, Fuchs G, Bacher A, Eisenreich W (2002). "A bicyclic autotrophic CO2 fixation pathway in Chloroflexus aurantiacus". J. Biol. Chem. 277 (23): 20277–83. PMID 11929869. doi:10.1074/jbc.M201030200.
Jump up ^ Zarzycki J, Brecht V, Müller M, Fuchs G (2009). "Identifying the missing steps of the autotrophic 3-hydroxypropionate CO2 fixation cycle in Chloroflexus aurantiacus". Proc. Natl. Acad. Sci. U.S.A. 106 (50): 21317–22. Bibcode:2009PNAS..10621317Z. PMC 2795484  . PMID 19955419. doi:10.1073/pnas.0908356106.
Jump up ^ Berg IA, Kockelkorn D, Buckel W, Fuchs G (2007). "A 3-hydroxypropionate/4-hydroxybutyrate autotrophic carbon dioxide assimilation pathway in Archaea". Science. 318 (5857): 1782–6. Bibcode:2007Sci...318.1782B. PMID 18079405. doi:10.1126/science.1149976.
Jump up ^ Huber H, Gallenberger M, Jahn U, Eylert E, Berg IA, Kockelkorn D, Eisenreich W, Fuchs G (2008). "A dicarboxylate/4-hydroxybutyrate autotrophic carbon assimilation cycle in the hyperthermophilic Archaeum Ignicoccus hospitalis". Proc. Natl. Acad. Sci. U.S.A. 105 (22): 7851–6. Bibcode:2008PNAS..105.7851H. PMC 2409403  . PMID 18511565. doi:10.1073/pnas.0801043105.
Jump up ^ Encyclopedia of Microbiology. Academic Press. 2009. pp. 83–84. ISBN 9780123739445.
Jump up ^ Nicole Kresge; Robert D. Simoni; Robert L. Hill (2005). "The Discovery of Heterotrophic Carbon Dioxide Fixation by Harland G. Wood". The Journal of Biological Chemistry.
Further reading[edit]
Berg IA (2011). "Ecological aspects of the distribution of different autotrophic CO2 fixation pathways". Appl. Environ. Microbiol. 77 (6): 1925–36. PMC 3067309  . PMID 21216907. doi:10.1128/AEM.02473-10.
Descent of plants and algae[edit]
Keeling PJ (2004). "Diversity and evolutionary history of plastids and their hosts". Am. J. Bot. 91 (10): 1481–93. PMID 21652304. doi:10.3732/ajb.91.10.1481.
Keeling PJ (2009). "Chromalveolates and the evolution of plastids by secondary endosymbiosis" (PDF). J. Eukaryot. Microbiol. 56 (1): 1–8. PMID 19335769. doi:10.1111/j.1550-7408.2008.00371.x. Retrieved 10 April 2012. [permanent dead link]
Keeling PJ (2010). "The endosymbiotic origin, diversification and fate of plastids". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 365 (1541): 729–48. PMC 2817223  . PMID 20124341. doi:10.1098/rstb.2009.0103.
Timme RE, Bachvaroff TR, Delwiche CF (2012). "Broad phylogenomic sampling and the sister lineage of land plants". PLoS ONE. 7 (1): e29696. Bibcode:2012PLoSO...7E9696T. PMC 3258253  . PMID 22253761. doi:10.1371/journal.pone.0029696.
Spiegel FW (2012). "Evolution. Contemplating the first Plantae". Science. 335 (6070): 809–10. Bibcode:2012Sci...335..809S. PMID 22344435. doi:10.1126/science.1218515.
Price DC, Chan CX, Yoon HS, Yang EC, Qiu H, Weber AP, Schwacke R, Gross J, Blouin NA, Lane C, Reyes-Prieto A, Durnford DG, Neilson JA, Lang BF, Burger G, Steiner JM, Löffelhardt W, Meuser JE, Posewitz MC, Ball S, Arias MC, Henrissat B, Coutinho PM, Rensing SA, Symeonidi A, Doddapaneni H, Green BR, Rajah VD, Boore J, Bhattacharya D (2012). "Cyanophora paradoxa genome elucidates origin of photosynthesis in algae and plants" (PDF). Science. 335 (6070): 843–7. Bibcode:2012Sci...335..843P. PMID 22344442. doi:10.1126/science.1213561. Retrieved 10 April 2012.
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text:
Patient Self-Determination Act
From Wikipedia, the free encyclopedia
Jump to: navigation, search
The Patient Self-Determination Act (PSDA) was passed by the United States Congress in 1990 as an amendment to the Omnibus Budget Reconciliation Act of 1990. Effective on December 1, 1991, this legislation required many hospitals, nursing homes, home health agencies, hospice providers, health maintenance organizations (HMOs), and other health care institutions to provide information about advance health care directives to adult patients upon their admission to the healthcare facility.[1][2] This law does not apply to individual physicians.
Because the 1991 PSDA does not apply to individual physicians, private clinics and practices, most of which are incorporated for-profit organizations, the problem of cruel over treatment for profit of the elderly on Medicare/Medicaid was not controlled to the extent that Congress hoped would be possible when the states would implement the goals of the 1991 PSDA in state laws. (Recently, in 2012, the newspapers have revealed that The United States Department of Health and Human Services and the Department of Justice have cooperated to prosecute over treatment of patients under the federal False Claims Act.) The goals of the 1991 PSDA are covered in the PURPOSE section, below.
Section 1233 of the proposed America's Affordable Health Choices Act of 2009 (H.R. 3200) would have authorized reimbursements for physician counseling regarding advance directives (once every five years)[3] but it was not included in the Patient Protection and Affordable Care Act of 2010 because of controversy over what were characterized as "death panels."[4][5]
Unfortunately, because the law of the 1991 PSDA does not require or mandate the treating physicians—after an educated "terminal" prognosis that is shared with the patient—to seek informed consent from elderly and terminal Medicare/Medicaid patients for either Curative Care under Medicare or palliative care, transition to Hospice in the last six months of life (also paid or out of the Medicare purse), the goals of the PSDA are not realized. Patient self rationing of expensive medical care at the end of life through the process of the advance directive under the provisions of the 1991 PSDA is discouraged because the patients have not had end-of-life conversations with the treating physicians.
Contents
[hide]
1 Requirements
2 Purpose
3 References
4 Further reading
5 External links
Requirements[edit]
The requirements of the PSDA are as follows:
Patients are given written notice upon admission to the health care facility of their decision-making rights, and policies regarding advance health care directives in their state and in the institution to which they have been admitted. Patient rights include:
The right to facilitate their own health care decisions
The right to accept or refuse medical treatment
The right to make an advance health care directive
Facilities must inquire as to whether the patient already has an advance health care directive, and make note of this in their medical records.
Facilities must provide education to their staff and affiliates about advance health care directives.
Health care providers are not allowed to discriminately admit or treat patients based on whether or not they have an advance health care directive.
Purpose[edit]
The purpose of the Patient Self-Determination Act was/is to inform patients of their rights regarding decisions toward their own medical care, and ensure that these rights are communicated by the health care provider. Specifically, the rights ensured are those of the patient to dictate their future care (by means such as living will or power of attorney), should they become incapacitated.
The goals of the "purpose" of the Patient Self-Determination Act are/were (1) to prevent cruel over treatment of elderly/disabled Medicare/Medicaid patients for the profit motive and (2) to save money for Medicare and the private insurers in the form of the reduction of end-of-life costs for Medicare and the private insurers when elderly Medicare/Medicaid patients would ELECT/CHOOSE to refuse expensive ICU/CCU life-extending or life-saving treatments in the hospital in order to shorten their suffering unto a certain death. It was envisioned by the framers of the 1991 PSDA that the elderly/disabled on Medicare/Medicaid would die less expensively and more comfortably (out of pain) on the palliative care/Hospice Medicare Entitlement in their own personal residence or in the setting of a residential nursing home when the treating physicians consulted with them about their terminal diagnoses.
References[edit]
Jump up ^ Health Care Advance Directives - What is the Patient Self-Determination Act?. American Bar Association.
Jump up ^ What is the Patient Self-Determination Act?. Legal HelpMate.
Jump up ^ Advance Care Planning in Health Care Reform Legislation Archived 2010-07-13 at the Wayback Machine.. National Hospice and Paliative Care Organization.
Jump up ^ Robert Pear (December 25, 2010). "Obama Returns to End-of-Life Plan That Caused Stir". The New York Times.
Jump up ^ Robert Pear (January 4, 2011). "U.S. Alters Rule on Paying for End-of-Life Planning". The New York Times.
Further reading[edit]
Kring, Daria L. (October–December 2007). "The Patient Self-determination Act: has it reached the end of its life?" . JONA'S Healthcare Law, Ethics and Regulation. 9 (4): 125–33. doi:10.1097/01.NHL.0000300767.91800.17. PMID 18043329.
Leahman, Dee (July–August 2004). "Why the Patient Self-Determination Act has failed". North Carolina Medical Journal. 65 (4): 249–51. PMID 15481498.
Pope, Thaddeus Mason (Winter 1999). "The Maladaptation of Miranda to Advance Directives: A Critique of the Patient Self Determination Act". Health Matrix. 9 (1): 139–202. PMID 10538190. SSRN 174749  .
Ulrich, Lawrence P. (1998), "The Requirements of the Patient Self-Determination", The Patient Self-Determination Act: Meeting the Challenges in Patient Care
Walerius, Theresa; Hill, Pamela D.; Anderson, Mary Ann (November–December 2009). "Nurses' knowledge of Advance Directives, Patient Self-determination Act, and Illinois Advance Directive Law". Clinical Nurse Specialist. 23 (6): 316–20. doi:10.1097/NUR.0b013e3181be3273. PMID 19858904.
Yates, Jeffery L.; Glick, Henry R. (1997). "The failed Patient Self-Determination Act and policy alternatives for the right to die" . Journal of Aging & Social Policy. 9 (4): 29–50. doi:10.1300/J031v09n04_03. PMID 10186890.
External links[edit]
"The Patient Self-Determination Act (PSDA)". American Cancer Society: Treatment Topics & Resources. June 15, 2009. Archived from the original on February 22, 2010.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Patient_Self-Determination_Act&oldid=843421267"
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