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'''Chlorophyll fluorescence''' is light that has been re-emitted after being absorbed by [[chlorophyll]] molecules of [[plant]] [[leaves]]. By measuring the intensity and nature of this [[fluorescence]], plant [[ecophysiology]] can be investigated. | |||
== Assessing plant physiology with Chlorophyll fluorescence == | |||
Light energy that has been absorbed by a leaf will excite [[electrons]] in [[chlorophyll]] molecules. Energy in [[photosystem II]] can be converted to [[chemical]] energy to drive [[photosynthesis]] (photochemistry). If photochemistry is inefficient, excess energy can damage the leaf. Energy can be emitted (known as energy quenching) in the form of heat (called [[non-photochemical quenching]]) or emitted as chlorophyll fluorescence. These three processes are in competition, so fluorescence yield is high when less energy is emitted as heat or used in photochemistry. Therefore, by measuring the amount of chlorophyll fluorescence, the efficiency of photochemistry and non-photochemical quenching can be assessed. | |||
The fluorescence emitted from a leaf has a longer wavelength than the light absorbed by the leaf. Therefore, fluorescence can be measured by shining a defined wavelength of light onto a leaf and measuring the level of light emitted at longer wavelengths. | |||
By measuring chlorophyll fluorescence, plant [[ecophysiology]] can be investigated. [[Chlorophyll fluorescence#Chlorophyll fluorometers|Chlorophyll fluorometers]] are used by plant researchers to assess plant stress. Gitelson (1999) states, "The ratio between chlorophyll fluorescence at 735 nm and the | |||
wavelength range 700nm to 710 nm, F735/F700 was found to be linearly proportional to the | |||
chlorophyll content (with determination coefficient, r2, more than 0.95) and thus this ratio can be used as a precise indicator of chlorophyll content in plant leaves."<ref>{{cite journal | doi = 10.1016/S0034-4257(99)00023-1 | title = The Chlorophyll Fluorescence Ratio F735/F700 as an Accurate Measure of the Chlorophyll Content in Plants | year = 1999 | last1 = Gitelson | first1 = Anatoly A | last2 = Buschmann | first2 = Claus | last3 = Lichtenthaler | first3 = Hartmut K | journal = Remote Sensing of Environment | volume = 69 | issue = 3 | pages = 296}}</ref> | |||
== The Kautsky effect == | |||
{{main|Kautsky effect}} | |||
Upon illumination of a dark-adapted leaf, there is a rapid rise in fluorescence from [[Photosystem II]] (PSII), followed by a slow decline. First observed by ''Kautsky et al., 1960'', this is called the Kautsky Effect. | |||
The increase in fluorescence is due to PSII [[reaction center]]s being in a "closed" state. Reaction centers are "closed" when unable to accept further electrons. This occurs when [[electron acceptor]]s downstream of PSII have not yet passed their electrons to a subsequent electron carrier, so are unable to accept another electron. Closed reaction centres reduce the overall photochemical efficiency, and so increases the level of fluorescence. Transferring a leaf from dark into light increases the proportion of closed PSII reaction centres, so fluorescence levels increase for 1–2 seconds. Subsequently, fluorescence decreases over a few minutes. This is due to; 1. more "photochemical quenching" in which electrons are transported away from PSII due to enzymes involved in carbon fixation; and 2. more "non-photochemical quenching" in which more energy is converted to heat. | |||
== Measuring fluorescence == | |||
Usually the initial measurement is the minimal level of fluorescence, <math>\,F_0</math>. This is the fluorescence in the absence of photosynthetic light.<ref name=guide/> | |||
To use measurements of chlorophyll fluorescence to analyse photosynthesis, researchers must distinguish between [[photochemical]] quenching and [[non-photochemical quenching]] (heat dissipation). This is achieved by stopping photochemistry, which allows researchers to measure fluorescence in the presence of non-photochemical quenching alone. To reduce photochemical quenching to negligible levels, a high intensity, short flash of light is applied to the leaf. This transiently closes all PSII reaction centres, which prevents energy of PSII being passed to downstream electron carriers. Non-photochemical quenching will not be effected if the flash is short. During the flash, the fluorescence reaches the level reached in the absence of any photochemical quenching, known as maximum fluorescence <math>\,F_m</math>.<ref name=guide>{{cite web|url=http://jxb.oxfordjournals.org/content/51/345/659.full.pdf+html |title=Chlorophyll fluorescence—a practical guide |publisher=Jxb.oxfordjournals.org |date=2000-04-01 |accessdate=2011-03-28}}</ref> | |||
The efficiency of photochemical quenching (which is a proxy of the efficiency of PSII) can be estimated by comparing <math>\,F_m</math> to the steady yield of fluorescence in the light <math>\,F_t</math> and the yield of fluorescence in the absence of photosynthetic light <math>\,F_0</math>.<ref name=opti/> | |||
The efficiency of non-photochemical quenching is altered by various internal and external factors. Alterations in heat dissipation mean changes in <math>\,F_m</math>. Heat dissipation cannot be totally stopped, so the yield of chlorophyll fluorescence in the absence of non-photochemical quenching cannot be measured. Therefore, researchers use a dark-adapted point (<math>F_m^0</math>) with which to compare estimations of non-photochemical quenching.<ref name=guide/><ref name=opti/> | |||
=== Common fluorescence parameters === | |||
<math>\,F_0</math>: Minimal fluorescence (arbitrary units). Fluorescence level when all antenna pigment complexes associated with the photosystem are assumed to be open (dark adapted).<ref name=opti>{{cite web|url=http://www.optisci.com/cf.htm |title=Stress Testing |publisher=Optisci.com |date= |accessdate=2011-03-28}}</ref> | |||
<math>\,F_m</math>: Maximal fluorescence (arbitrary units). Fluorescence level when a high intensity flash has been applied. All antenna sites are assumed to be closed.<ref name=opti/> | |||
<math>\,F_{tr}</math>: Terminal fluorescence (arbitrary units). Fluorescence quenching value at the end of the test. | |||
<math>\,T_{1/2}</math>: Half rise time from <math>\,F_0</math> to <math>\,F_m</math>. | |||
=== Calculated parameters === | |||
<math>\,F_v</math> is variable fluorescence. Calculated as <math>\,F_v</math> = <math>\,F_m</math> - <math>\,F_0</math>.<ref name=neo/> | |||
<math>\tfrac{F_v}{F_m}</math> is the ratio of variable fluorescence to maximal fluorescence. | |||
Calculated as <math>\frac{F_m-F_0}{F_m}</math>. This is a measure of the maximum efficiency of PSII (the efficiency if all PSII centres were open). <math>\tfrac{F_v}{F_m}</math> can be used to estimate the potential efficiency of PSII by taking dark-adapted measurements.<ref name=opti/> | |||
<math>\,\Phi_{PSII}</math> measures the efficiency of Photosystem II. Calculated as <math>\frac{F_m-F_{tr}}{F_m}</math>. This parameter measures the proportion of light absorbed by PSII that is used in photochemistry. As such, it can give a measure of the rate of linear electron transport and so indicates overall photosynthesis. | |||
<math>\,qP</math> (photochemical quenching). Calculated as <math>\,\frac{F_m-F_{tr}}{F_m-F_0}</math>. This parameter approximates the proportion of PSII reaction centres that are open. | |||
Whilst <math>\,\Phi_{PSII}</math> gives an estimation of the efficiency, <math>\,qP</math> and <math>\tfrac{F_v}{F_m}</math> tell us which processes which have altered the efficiency. Closure of reaction centers as a result of a high intensity light will alter the value of <math>\,qP</math>. Changes in the efficiency of non-photochemical quenching will alter the ratio <math>\tfrac{F_v}{F_m}</math>.<ref name=opti/> | |||
== Applications of the Theory == | |||
=== PSII yield as a measure of photosynthesis === | |||
Chlorophyll fluorescence appears to measure of photosynthesis, but this is an over-simplification. Fluorescence can measure the efficiency of PSII photochemistry, which can be used to estimate the rate of linear electron transport by multiplying by the light intensity. However, researchers generally mean [[carbon fixation]] when they refer to photosynthesis. Electron transport and CO<sub>2</sub> fixation can correlate well, but may not correlate in the field due to processes such as photorespiration, nitrogen metabolism and the [[Mehler reaction]]. | |||
* De Martino ''et al.'' (2007)<ref>{{cite web|url=http://www.springerlink.com/content/j210g33838876662/ |title=Abstract:Preliminary investigation into the uneven ripening of banana peel after 1-MCP treatment | |||
|publisher=SpringerLink |year=2007 |author=De Martino et al. |doi=10.1007/978-1-4020-6014-4_53 | |||
}}</ref> examined [[banana peel]] degreening after treatment with [[1-methylcyclopropene]] (1-MCP). Many parameters were measured, including chlorophyll fluorescence since fluorescense is a measure of [[chloroplast]] photochemical efficiency. 1-MCP treatment was shown to delay the decrease in chlorophyll fluorescence. Greenness and photochemical efficiency decreased simultaneously after ethylene treatment, suggesting that changes in chlorophyll fluorescence <math>\,F_m</math> were responsible for the patterns of de-greening. After 7 and 8 days, <math>\,F_m</math> values of 1-MCP peel were ca. 3-fold the values of the other treatments, showing that 1-MCP treated fruit maintain their photochemical efficiency when the other treatment regimes do not. | |||
=== Relating electron transport to carbon fixation === | |||
A powerful research technique is to simultaneously measure chlorophyll fluorescence and [[gas exchange]] to obtain a full picture of the response of plants to their environment. One technique is to simultaneously measure CO<sub>2</sub> fixation and PSII photochemistry at different light intensities, in non-photorespiratory conditions. A plot of CO<sub>2</sub> fixation and PSII photochemistry indicates the electron requirement per molecule CO<sub>2</sub> fixed. From this estimation, the extent of [[photorespiration]] may be estimated. This has been used to explore the significance of photorespiration as a photoprotective mechanism during drought. | |||
Fluorescence analysis can also be applied to understanding the effects of low and high temperatures. | |||
* Sobrado (2008)<ref>{{cite web|url=http://www.springerlink.com/content/d14258646q4pq146/fulltext.pdf |title=Leaf characteristics and diurnal variation of chlorophyll fluorescence in leaves of the ‘bana’ vegetation of the amazon region |format=PDF |year=2008 |author=Sobrado}}</ref> investigated gas exchange and chlorophyll ''a'' fluorescence responses to high intensity light, of pioneer species and forest species. Midday leaf gas exchange was measured using a [[photosynthesis system]], which measured net photosynthetic rate, gs, and intercellular CO<sub>2</sub> concentration (<math>C_i</math>). In the same leaves used for gas exchange measurements, chlorophyll ''a'' fluorescence parameters (initial, <math>\,F_0</math>; maximum, <math>\,F_m</math>; and variable, <math>\,F_v</math>) were measured using a fluorometer. The results showed that despite pioneer species and forest species occupying different habitats, both showed similar vulnerability to midday photoinhibition in sun-exposed leaves. | |||
=== Measuring stress and stress tolerance === | |||
Chlorophyll fluorescence can measure most types of [[plant stress measurement|plant stress]]. Chlorophyll fluorescence can be used as a proxy of plant stress because environmental stresses, e.g. extremes of temperature, light and water availability, can reduce the ability of a plant to metabolise normally. This can mean an imbalance between the absorption of light energy by chlorophyll and the use of energy in photosynthesis.<ref>{{cite web|url=http://personalpages.manchester.ac.uk/staff/giles.johnson/default.php?page=research |title=Plant Stress Biology |publisher=Personalpages.manchester.ac.uk |date= |accessdate=2011-03-28}}</ref> | |||
* Favaretto et al. (2010)<ref>{{cite journal |url=http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6T66-50CV80C-2&_user=10&_coverDate=01%2F31%2F2011&_rdoc=1&_fmt=high&_orig=search&_origin=search&_sort=d&_docanchor=&view=c&_searchStrId=1586619461&_rerunOrigin=google&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=0fa9176d9f11b6ab2b20cdfefcc5697a&searchtype=a |title=Differential responses of antioxidant enzymes in pioneer and late-successional tropical tree species grown under sun and shade conditions |author=Favaretto et al. |year=2011 | |||
}}</ref> investigated adaptation to a strong light environment in pioneer and late successional species, grown under 100% and 10% light. Numerous parameters, including chlorophyll ''a'' fluorescence, were measured. A greater decline in <math>\tfrac{F_v}{F_m}</math> under full sun light in the late-successional species than in the pioneer species was observed. Overall, their results show that pioneer species perform better under high-sun light than late- successional species, suggesting that pioneer plants have more potential tolerance to photo-oxidative damage. | |||
* Neocleous and Vasilakakis (2009)<ref name=neo>{{cite web|url=http://www.informaworld.com/smpp/content~db=all~content=a906673051~frm=abslink |title=Effects of Boron and Salinity on Red Raspberry in Vitro - International Journal of Fruit Science |publisher=Informaworld.com |date=2008-12-03 |accessdate=2011-03-28}}</ref> investigated the response of [[raspberry]] to [[boron]] and [[salt]] stress. An chlorophyll fluorometer was used to measure <math>\,F_0</math>, <math>\,F_m</math> and <math>\,F_v</math>. The leaf chlorophyll fluorescence was not significantly affected by NaCl concentration when B concentration was low. When B was increased, leaf chlorophyll fluorescence was reduced under saline conditions. It could be concluded that the combined effect of B and NaCl on raspberries induces a toxic effect in photochemical parameters. | |||
== Chlorophyll fluorometers == | |||
The development of portable [[fluorometer]]s has allowed chlorophyll fluorescence to become a common method of measuring plant stress in plant ecophysiology studies. Chlorophyll fluorescence has been revolutionized by the use of modulated chlorophyll fluorometers in which the light source is modulated (rapidly switched on and off) and the detector is tuned to detect only fluorescence excited by the measuring light. This means the relative yield of fluorescence can be measured in the presence of background light. Crucially, this means chlorophyll fluorescence can be measured in the field in full sunlight.<ref name=guide/> Most modern fluorometers are modulated.[[File:CCM300 being used to measure the chlorophyll content of small samples.jpg|thumb|By measuring chlorophyll content nitrogen deficiency can be detected.]] | |||
Some modulated fluorometers can determine both ambient light and dark adaptation parameters (Fo, Fm, Fv/Fm, Y, Ft, Foq, Fms and OJIP transients) and can calculate photochemical and non-photochemical quenching coefficients (qP, qN and NPQ). In contrast, some fluorometers are slimmed down, and designed to be portable and operated in one hand. | |||
A recent advancement in chlorophyll fluorescence is the development of imaging fluorometers which can visualize spatial heterogeneities in photosynthetic activity of a sample. | |||
Because of the link between chlorophyll content and [[nitrogen]] content in leaves, chlorophyll fluorometers can be used to detect nitrogen deficiency in plants, by [[Plant tissue test#Chlorophyll fluorometry|several methods]]. | |||
== See also == | |||
[[non-photochemical quenching]] | |||
== References == | |||
<references /> | |||
==External links== | |||
* Kalaji et al. 2012. Experimental in vivo measurements of light emission in plants: a perspective dedicated to David Walker, http://link.springer.com/article/10.1007%2Fs11120-012-9780-3 | |||
*{{cite journal | doi = 10.1093/jexbot/51.345.659 | title = Chlorophyll fluorescence--a practical guide | year = 2000 | last1 = Maxwell | first1 = K. | journal = Journal of Experimental Botany | volume = 51 | issue = 345 | pages = 659–68 | pmid = 10938857 | last2 = Johnson | first2 = GN}} | |||
{{DEFAULTSORT:Chlorophyll Fluorescence}} | |||
[[Category:Light reactions]] |
Revision as of 12:04, 21 October 2013
Chlorophyll fluorescence is light that has been re-emitted after being absorbed by chlorophyll molecules of plant leaves. By measuring the intensity and nature of this fluorescence, plant ecophysiology can be investigated.
Assessing plant physiology with Chlorophyll fluorescence
Light energy that has been absorbed by a leaf will excite electrons in chlorophyll molecules. Energy in photosystem II can be converted to chemical energy to drive photosynthesis (photochemistry). If photochemistry is inefficient, excess energy can damage the leaf. Energy can be emitted (known as energy quenching) in the form of heat (called non-photochemical quenching) or emitted as chlorophyll fluorescence. These three processes are in competition, so fluorescence yield is high when less energy is emitted as heat or used in photochemistry. Therefore, by measuring the amount of chlorophyll fluorescence, the efficiency of photochemistry and non-photochemical quenching can be assessed.
The fluorescence emitted from a leaf has a longer wavelength than the light absorbed by the leaf. Therefore, fluorescence can be measured by shining a defined wavelength of light onto a leaf and measuring the level of light emitted at longer wavelengths.
By measuring chlorophyll fluorescence, plant ecophysiology can be investigated. Chlorophyll fluorometers are used by plant researchers to assess plant stress. Gitelson (1999) states, "The ratio between chlorophyll fluorescence at 735 nm and the wavelength range 700nm to 710 nm, F735/F700 was found to be linearly proportional to the chlorophyll content (with determination coefficient, r2, more than 0.95) and thus this ratio can be used as a precise indicator of chlorophyll content in plant leaves."[1]
The Kautsky effect
Mining Engineer (Excluding Oil ) Truman from Alma, loves to spend time knotting, largest property developers in singapore developers in singapore and stamp collecting. Recently had a family visit to Urnes Stave Church. Upon illumination of a dark-adapted leaf, there is a rapid rise in fluorescence from Photosystem II (PSII), followed by a slow decline. First observed by Kautsky et al., 1960, this is called the Kautsky Effect.
The increase in fluorescence is due to PSII reaction centers being in a "closed" state. Reaction centers are "closed" when unable to accept further electrons. This occurs when electron acceptors downstream of PSII have not yet passed their electrons to a subsequent electron carrier, so are unable to accept another electron. Closed reaction centres reduce the overall photochemical efficiency, and so increases the level of fluorescence. Transferring a leaf from dark into light increases the proportion of closed PSII reaction centres, so fluorescence levels increase for 1–2 seconds. Subsequently, fluorescence decreases over a few minutes. This is due to; 1. more "photochemical quenching" in which electrons are transported away from PSII due to enzymes involved in carbon fixation; and 2. more "non-photochemical quenching" in which more energy is converted to heat.
Measuring fluorescence
Usually the initial measurement is the minimal level of fluorescence, . This is the fluorescence in the absence of photosynthetic light.[2]
To use measurements of chlorophyll fluorescence to analyse photosynthesis, researchers must distinguish between photochemical quenching and non-photochemical quenching (heat dissipation). This is achieved by stopping photochemistry, which allows researchers to measure fluorescence in the presence of non-photochemical quenching alone. To reduce photochemical quenching to negligible levels, a high intensity, short flash of light is applied to the leaf. This transiently closes all PSII reaction centres, which prevents energy of PSII being passed to downstream electron carriers. Non-photochemical quenching will not be effected if the flash is short. During the flash, the fluorescence reaches the level reached in the absence of any photochemical quenching, known as maximum fluorescence .[2]
The efficiency of photochemical quenching (which is a proxy of the efficiency of PSII) can be estimated by comparing to the steady yield of fluorescence in the light and the yield of fluorescence in the absence of photosynthetic light .[3] The efficiency of non-photochemical quenching is altered by various internal and external factors. Alterations in heat dissipation mean changes in . Heat dissipation cannot be totally stopped, so the yield of chlorophyll fluorescence in the absence of non-photochemical quenching cannot be measured. Therefore, researchers use a dark-adapted point () with which to compare estimations of non-photochemical quenching.[2][3]
Common fluorescence parameters
: Minimal fluorescence (arbitrary units). Fluorescence level when all antenna pigment complexes associated with the photosystem are assumed to be open (dark adapted).[3]
: Maximal fluorescence (arbitrary units). Fluorescence level when a high intensity flash has been applied. All antenna sites are assumed to be closed.[3]
: Terminal fluorescence (arbitrary units). Fluorescence quenching value at the end of the test.
Calculated parameters
is variable fluorescence. Calculated as = - .[4]
is the ratio of variable fluorescence to maximal fluorescence. Calculated as . This is a measure of the maximum efficiency of PSII (the efficiency if all PSII centres were open). can be used to estimate the potential efficiency of PSII by taking dark-adapted measurements.[3]
measures the efficiency of Photosystem II. Calculated as . This parameter measures the proportion of light absorbed by PSII that is used in photochemistry. As such, it can give a measure of the rate of linear electron transport and so indicates overall photosynthesis.
(photochemical quenching). Calculated as . This parameter approximates the proportion of PSII reaction centres that are open.
Whilst gives an estimation of the efficiency, and tell us which processes which have altered the efficiency. Closure of reaction centers as a result of a high intensity light will alter the value of . Changes in the efficiency of non-photochemical quenching will alter the ratio .[3]
Applications of the Theory
PSII yield as a measure of photosynthesis
Chlorophyll fluorescence appears to measure of photosynthesis, but this is an over-simplification. Fluorescence can measure the efficiency of PSII photochemistry, which can be used to estimate the rate of linear electron transport by multiplying by the light intensity. However, researchers generally mean carbon fixation when they refer to photosynthesis. Electron transport and CO2 fixation can correlate well, but may not correlate in the field due to processes such as photorespiration, nitrogen metabolism and the Mehler reaction.
- De Martino et al. (2007)[5] examined banana peel degreening after treatment with 1-methylcyclopropene (1-MCP). Many parameters were measured, including chlorophyll fluorescence since fluorescense is a measure of chloroplast photochemical efficiency. 1-MCP treatment was shown to delay the decrease in chlorophyll fluorescence. Greenness and photochemical efficiency decreased simultaneously after ethylene treatment, suggesting that changes in chlorophyll fluorescence were responsible for the patterns of de-greening. After 7 and 8 days, values of 1-MCP peel were ca. 3-fold the values of the other treatments, showing that 1-MCP treated fruit maintain their photochemical efficiency when the other treatment regimes do not.
Relating electron transport to carbon fixation
A powerful research technique is to simultaneously measure chlorophyll fluorescence and gas exchange to obtain a full picture of the response of plants to their environment. One technique is to simultaneously measure CO2 fixation and PSII photochemistry at different light intensities, in non-photorespiratory conditions. A plot of CO2 fixation and PSII photochemistry indicates the electron requirement per molecule CO2 fixed. From this estimation, the extent of photorespiration may be estimated. This has been used to explore the significance of photorespiration as a photoprotective mechanism during drought.
Fluorescence analysis can also be applied to understanding the effects of low and high temperatures.
- Sobrado (2008)[6] investigated gas exchange and chlorophyll a fluorescence responses to high intensity light, of pioneer species and forest species. Midday leaf gas exchange was measured using a photosynthesis system, which measured net photosynthetic rate, gs, and intercellular CO2 concentration (). In the same leaves used for gas exchange measurements, chlorophyll a fluorescence parameters (initial, ; maximum, ; and variable, ) were measured using a fluorometer. The results showed that despite pioneer species and forest species occupying different habitats, both showed similar vulnerability to midday photoinhibition in sun-exposed leaves.
Measuring stress and stress tolerance
Chlorophyll fluorescence can measure most types of plant stress. Chlorophyll fluorescence can be used as a proxy of plant stress because environmental stresses, e.g. extremes of temperature, light and water availability, can reduce the ability of a plant to metabolise normally. This can mean an imbalance between the absorption of light energy by chlorophyll and the use of energy in photosynthesis.[7]
- Favaretto et al. (2010)[8] investigated adaptation to a strong light environment in pioneer and late successional species, grown under 100% and 10% light. Numerous parameters, including chlorophyll a fluorescence, were measured. A greater decline in under full sun light in the late-successional species than in the pioneer species was observed. Overall, their results show that pioneer species perform better under high-sun light than late- successional species, suggesting that pioneer plants have more potential tolerance to photo-oxidative damage.
- Neocleous and Vasilakakis (2009)[4] investigated the response of raspberry to boron and salt stress. An chlorophyll fluorometer was used to measure , and . The leaf chlorophyll fluorescence was not significantly affected by NaCl concentration when B concentration was low. When B was increased, leaf chlorophyll fluorescence was reduced under saline conditions. It could be concluded that the combined effect of B and NaCl on raspberries induces a toxic effect in photochemical parameters.
Chlorophyll fluorometers
The development of portable fluorometers has allowed chlorophyll fluorescence to become a common method of measuring plant stress in plant ecophysiology studies. Chlorophyll fluorescence has been revolutionized by the use of modulated chlorophyll fluorometers in which the light source is modulated (rapidly switched on and off) and the detector is tuned to detect only fluorescence excited by the measuring light. This means the relative yield of fluorescence can be measured in the presence of background light. Crucially, this means chlorophyll fluorescence can be measured in the field in full sunlight.[2] Most modern fluorometers are modulated.
Some modulated fluorometers can determine both ambient light and dark adaptation parameters (Fo, Fm, Fv/Fm, Y, Ft, Foq, Fms and OJIP transients) and can calculate photochemical and non-photochemical quenching coefficients (qP, qN and NPQ). In contrast, some fluorometers are slimmed down, and designed to be portable and operated in one hand.
A recent advancement in chlorophyll fluorescence is the development of imaging fluorometers which can visualize spatial heterogeneities in photosynthetic activity of a sample.
Because of the link between chlorophyll content and nitrogen content in leaves, chlorophyll fluorometers can be used to detect nitrogen deficiency in plants, by several methods.
See also
References
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- ↑ 3.0 3.1 3.2 3.3 3.4 3.5 Template:Cite web
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- ↑ Template:Cite web
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- ↑ Template:Cite web
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In its statement, the singapore property listing - website link, government claimed that the majority citizens buying their first residence won't be hurt by the new measures. Some concessions can even be prolonged to chose teams of consumers, similar to married couples with a minimum of one Singaporean partner who are purchasing their second property so long as they intend to promote their first residential property. Lower the LTV limit on housing loans granted by monetary establishments regulated by MAS from 70% to 60% for property purchasers who are individuals with a number of outstanding housing loans on the time of the brand new housing purchase. Singapore Property Measures - 30 August 2010 The most popular seek for the number of bedrooms in Singapore is 4, followed by 2 and three. Lush Acres EC @ Sengkang
Discover out more about real estate funding in the area, together with info on international funding incentives and property possession. Many Singaporeans have been investing in property across the causeway in recent years, attracted by comparatively low prices. However, those who need to exit their investments quickly are likely to face significant challenges when trying to sell their property – and could finally be stuck with a property they can't sell. Career improvement programmes, in-house valuation, auctions and administrative help, venture advertising and marketing, skilled talks and traisning are continuously planned for the sales associates to help them obtain better outcomes for his or her shoppers while at Knight Frank Singapore. No change Present Rules
Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.
A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running
The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more
There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang
External links
- Kalaji et al. 2012. Experimental in vivo measurements of light emission in plants: a perspective dedicated to David Walker, http://link.springer.com/article/10.1007%2Fs11120-012-9780-3
- One of the biggest reasons investing in a Singapore new launch is an effective things is as a result of it is doable to be lent massive quantities of money at very low interest rates that you should utilize to purchase it. Then, if property values continue to go up, then you'll get a really high return on funding (ROI). Simply make sure you purchase one of the higher properties, reminiscent of the ones at Fernvale the Riverbank or any Singapore landed property Get Earnings by means of Renting
In its statement, the singapore property listing - website link, government claimed that the majority citizens buying their first residence won't be hurt by the new measures. Some concessions can even be prolonged to chose teams of consumers, similar to married couples with a minimum of one Singaporean partner who are purchasing their second property so long as they intend to promote their first residential property. Lower the LTV limit on housing loans granted by monetary establishments regulated by MAS from 70% to 60% for property purchasers who are individuals with a number of outstanding housing loans on the time of the brand new housing purchase. Singapore Property Measures - 30 August 2010 The most popular seek for the number of bedrooms in Singapore is 4, followed by 2 and three. Lush Acres EC @ Sengkang
Discover out more about real estate funding in the area, together with info on international funding incentives and property possession. Many Singaporeans have been investing in property across the causeway in recent years, attracted by comparatively low prices. However, those who need to exit their investments quickly are likely to face significant challenges when trying to sell their property – and could finally be stuck with a property they can't sell. Career improvement programmes, in-house valuation, auctions and administrative help, venture advertising and marketing, skilled talks and traisning are continuously planned for the sales associates to help them obtain better outcomes for his or her shoppers while at Knight Frank Singapore. No change Present Rules
Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.
A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running
The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more
There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang