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New moon
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For other uses, see New moon (disambiguation).
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A graphic depicting the new moon phase
In astronomy, the new moon is the first lunar phase, when the Moon and Sun have the same ecliptic longitude.[1] At this phase, the lunar disk is not visible to the unaided eye, except when silhouetted during a solar eclipse. Daylight outshines the earthlight that dimly illuminates the dark side of the new Moon. The actual phase is usually a very thin crescent because the Moon rarely passes directly in front of the Sun, except in a solar eclipse.[note 1]
The original meaning of the term new moon, which is still sometimes used in non-astronomical contexts, was the first visible crescent of the Moon, after conjunction with the Sun.[2] This crescent Moon is briefly visible when low above the western horizon shortly after sunset and before moonset.
A lunation or synodic month is the average time from one new moon to the next. In the J2000.0 epoch, the average length of a lunation is 29.530588 days (or 29 days, 12 hours, 44 minutes, and 2.8 seconds). However, the length of any one synodic month can vary from 29.26 to 29.80 days due to the perturbing effects of the Sun's gravity on the Moon's eccentric orbit.[3] In a lunar calendar, each month corresponds to a lunation. Each lunar cycle can be assigned a unique lunation number to identify it.
Contents
1 Determining new moons: an approximate formula
1.1 Explanation of the formula
2 Lunation Number
3 Lunar calendars
3.1 Hindu calendar
3.2 Islamic calendar
3.3 Chinese calendar
3.4 Jewish calendar
3.5 Bahá'í calendar
4 See also
5 Notes
6 References
7 External links
Determining new moons: an approximate formula[edit]
This article contains instructions, advice, or how-to content. The purpose of Wikipedia is to present facts, not to train. Please help improve this article either by rewriting the how-to content or by moving it to Wikiversity, Wikibooks or Wikivoyage. (January 2018)
The length of a lunation is about 29.53 days. Its precise duration is linked to many phenomena in nature, such as the variation between spring and neap tides (the most and least profound tidal variances respectively). An approximate formula to compute the mean moments of new moon (conjunction between Sun and Moon) for successive months is:
d = 5.597661 + 29.5305888610 × N + ( 102.026 × 10 − 12 ) × N 2 {\displaystyle d=5.597661+29.5305888610\times N+(102.026\times 10^{-12})\times N^{2}}
where N is an integer, starting with 0 for the first new moon in the year 2000, and that is incremented by 1 for each successive synodic month; and the result d is the number of days (and fractions) since 2000-01-01 00:00:00 reckoned in the time scale known as Terrestrial Time (TT) used in ephemerides.
To obtain this moment expressed in Universal Time (UT, world clock time), add the result of following approximate correction to the result d obtained above:
− 0.000739 − ( 235 × 10 − 12 ) × N 2 {\displaystyle -0.000739-(235\times 10^{-12})\times N^{2}} days
Periodic perturbations change the time of true conjunction from these mean values. For all new moons between 1601 and 2401, the maximum difference is 0.592 days = 14h13m in either direction. The duration of a lunation (i.e. the time from new moon to the next new moon) varies in this period between 29.272 and 29.833 days, i.e. −0.259d = 6h12m shorter, or +0.302d = 7h15m longer than average.[4][5] This range is smaller than the difference between mean and true conjunction, because during one lunation the periodic terms cannot all change to their maximum opposite value.
See the article on the full moon cycle for a fairly simple method to compute the moment of new moon more accurately.
The long-term error of the formula is approximately: 1 cy2 seconds in TT, and 11 cy2 seconds in UT (cy is centuries since 2000; see section Explanation of the formulae for details.)
Explanation of the formula[edit]
The moment of mean conjunction can easily be computed from an expression for the mean ecliptical longitude of the Moon minus the mean ecliptical longitude of the Sun (Delauney parameter D). Jean Meeus gave formulae to compute this in his Astronomical Formulae for Calculators based on the ephemerides of Brown and Newcomb (ca. 1900); and in his 1st edition of Astronomical Algorithms[6] based on the ELP2000-85[7] (the 2nd edition uses ELP2000-82 with improved expressions from Chapront et al. in 1998). These are now outdated: Chapront et al. (2002)[8] published improved parameters. Also Meeus's formula uses a fractional variable to allow computation of the four main phases, and uses a second variable for the secular terms. For the convenience of the reader, the formula given above is based on Chapront's latest parameters and expressed with a single integer variable, and the following additional terms have been added:
constant term:
Like Meeus, apply the constant terms of the aberration of light for the Sun's motion and light-time correction for the Moon[note 2] to obtain the apparent difference in ecliptical longitudes:
Sun: +20.496"[9]
Moon: −0.704"[10]
Correction in conjunction: −0.000451 days[note 3]
For UT: at 1 January 2000, ΔT (= TT − UT ) was +63.83 s;[note 4] hence the correction for the clock time UT = TT − ΔT of the conjunction is:
−0.000739 days.
quadratic term:
In ELP2000–85 (see Chapront et alii 1988), D has a quadratic term of −5.8681"T2; expressed in lunations N, this yields a correction of +87.403×10–12N2[note 5] days to the time of conjunction. The term includes a tidal contribution of 0.5×(−23.8946 "/cy2). The most current estimate from Lunar Laser Ranging for the acceleration is (see Chapront et alii 2002): (−25.858 ±0.003)"/cy2. Therefore, the new quadratic term of D is = -6.8498"T2.[note 6] Indeed, the polynomial provided by Chapront et alii (2002) provides the same value (their Table 4). This translates to a correction of +14.622×10−12N2 days to the time of conjunction; the quadratic term now is:
+102.026×10−12N2 days.
For UT: analysis of historical observations shows that ΔT has a long-term increase of +31 s/cy2.[11] Converted to days and lunations,[note 7] the correction from ET to UT becomes:
−235×10−12N2 days.
The theoretical tidal contribution to ΔT is about +42 s/cy2[12] the smaller observed value is thought to be mostly due to changes in the shape of the Earth.[13] Because the discrepancy is not fully explained, uncertainty of our prediction of UT (rotation angle of the Earth) may be as large as the difference between these values: 11 s/cy2. The error in the position of the Moon itself is only maybe 0.5"/cy2,[note 8] or (because the apparent mean angular velocity of the Moon is about 0.5"/s), 1 s/cy2 in the time of conjunction with the Sun.
Lunation Number[edit]
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The Lunation Number or Lunation Cycle is a number given to each lunation beginning from a certain one in history. Several conventions are in use.[14]
The most commonly used is the Brown Lunation Number (BLN), which defines lunation 1 as beginning at the first new moon of 1923, the year when Ernest William Brown's lunar theory was introduced in the major national astronomical almanacs. Lunation 1 occurred at approximately 02:41 UTC, January 17, 1923. New moons occur on Julian Dates 2449128.59 + 29.53058867 ∗ ( B L N − 871 ) ± 0.25 {\displaystyle 2449128.59+29.53058867*(\mathrm {BLN} -871)\pm 0.25} , with the given uncertainty due to varying torques from the sun.
Another increasingly popular lunation number (simply called the Lunation Number), introduced by Jean Meeus, defines lunation 0 as beginning on the first new moon of 2000 (this occurred at approximately 18:14 UTC, January 6, 2000). The formula relating this Lunation Number with the Brown Lunation Number is: BLN = LN + 953.
The Goldstine Lunation Number refers to the lunation numbering used by Herman Goldstine in his 1973 book New and Full Moons: 1001 B.C. to A.D. 1651, with lunation 0 beginning on January 11, 1001 BC, and can be calculated using GLN = LN + 37105.
The Hebrew Lunation Number is the count of lunations in the Hebrew calendar with lunation 1 beginning on October 7, 3761 BC. It can be calculated using HLN = LN + 71234. The Islamic Lunation Number is the count of lunations in the Islamic calendar with lunation 1 as beginning on July 16, 622. It can be calculated using ILN = LN + 17038. The Thai Lunation Number is called "มาสเกณฑ์" (Maasa-Kendha), defines lunation 0 as beginning of the SouthEast-Asian Calendar on Sunday March 22, 638 (Julian Calendar). It can be calculated using TLN = LN + 16843.
Lunar calendars[edit]
See also: Lunar calendar
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In non-astronomical contexts, new moon refers to the first visible crescent of the Moon, after conjunction with the Sun.[15] This takes place over the western horizon in a brief period between sunset and moonset, and therefore the precise time and even the date of the appearance of the new moon by this definition will be influenced by the geographical location of the observer. The astronomical new moon, sometimes known as the dark moon to avoid confusion, occurs by definition at the moment of conjunction in ecliptical longitude with the Sun, when the Moon is invisible from the Earth. This moment is unique and does not depend on location, and in certain circumstances it coincides with a solar eclipse.
In the above meaning,[citation needed] the first crescent marks the beginning of the month in the Islamic calendar, and in some lunisolar calendars such as the Hebrew calendar. In the Chinese calendar, the beginning of the month is marked by the dark moon. The new moon is also important in astrology, as is the full moon.[citation needed]
Hindu calendar[edit]
The new moon is significant in the Hindu calendar. People[who?] generally wait for the new moon to begin projects, as the waxing period of the moon is considered to be favorable for new work.
There are fifteen moon dates for each of the waxing and waning periods. These fifteen dates divided evenly into five categories: Nanda, Bhadra', Jaya, Rikta, and Purna, which are cycled through in that order.
Nanda dates are considered to be favorable for auspicious works; Bhadra dates for works related with community, social, family, friends; and Jaya dates for dealing with conflict. Rikta dates are considered beneficial only for works related to cruelty. Purna dates are considered to be favorable for all work.
The first day of the Lunar Hindu calendar starts the day after the new moon (Amavasya), which is considered a powerful force for good or evil. The Hindu epic Mahabharatha states that the Kurukshetra War started this day, which was also a Tuesday (Mangalvaar, day of the week named after Mars).[citation needed]
Islamic calendar[edit]
The Islamic calendar has retained an observational definition of the new moon, marking the new month when the first crescent moon is actually seen, and making it impossible to be certain in advance of when a specific month will begin (in particular, the exact date on which Ramadan will begin is not known in advance). In Saudi Arabia, the new King Abdullah Centre for Crescent Observations and Astronomy in Mecca has a clock for addressing this as an international scientific project.[citation needed] In Pakistan, there is a "Central Ruet-e-Hilal Committee" whose head is Mufti Muneeb-ur-Rehman, assisted by 150 observatories of the Pakistan Meteorological Department, which announces the sighting of the new moon. Since its creation in 1974, the status of the Central Ruet-e-Hilal Committee has been controversial as it refused the "Witnesses" (Shahadats) from other sects.[16]
An attempt to unify Muslims on a scientifically calculated worldwide calendar was adopted by both the Fiqh Council of North America and the European Council for Fatwa and Research in 2007. The new calculation requires that conjunction must occur before sunset in Mecca, Saudi Arabia, and that, in the same evening, moonset must take place after sunset. These can be precisely calculated and therefore a unified calendar is imminent if it becomes adopted worldwide.[17][18]
Chinese calendar[edit]
The new moon is the beginning of the month in the Chinese calendar. Some Buddhist Chinese keep a vegetarian diet on the new moon and full moon each month.[19]
Jewish calendar[edit]
Main article: Rosh Chodesh
The new moon signifies the start of every Jewish month, and is considered an important date and minor holiday in the Hebrew calendar. The modern form of the calendar is a rule-based lunisolar calendar, akin to the Chinese calendar, measuring months defined in lunar cycles as well as years measured in solar cycles, and distinct from the purely lunar Islamic calendar and the predominantly solar Gregorian calendar. The Jewish calendar is calculated based on mathematical rules designed to ensure that festivals are observed in their traditional season. Passover always falls in the springtime.[20] This fixed lunisolar calendar follows rules introduced by Hillel II and refined until the ninth century This calculation makes use of a mean lunation length used by Ptolemy and handed down from Babylonians, which is still very accurate: ca. 29.530594 days vs. a present value (see below) of 29.530589 days. This difference of only 0.0000005, or five millionths of a day, adds up to about only four hours since Babylonian times.[citation needed]
The messianic Pentecostal group, the New Israelites of Peru, keeps the new moon as a Sabbath of rest. As an evangelical church, it follows the Bible's teachings that God sanctified the seventh-day Sabbath, and the new moons in addition to it. See Ezekiel 46:1, 3. No work may be done from dusk until dusk, and the services run for 11 hours, although a large number spend 24 hours within the gates of the temples, sleeping and singing praises throughout the night.[citation needed]
Bahá'í calendar[edit]
In the Bahá'í Faith, effective from 2015 onwards, the "Twin Holy Birthdays", referring to two successive holy days in the Bahá'í calendar (the birth of the Báb and the birth of Bahá'u'lláh), will be observed on the first and the second day following the occurrence of the eighth new moon after Naw-Rúz (Bahá'í New Year), as determined in advance by astronomical tables using Tehran as the point of reference.[21] This will result in the observance of the Twin Birthdays moving, year to year, from mid-October to mid-November according to the Gregorian calendar.[22]
See also[edit]
Moon portal
Solar eclipse
Dark moon
Black moon
Blue moon
Hilal
Lunar phase
Wet moon
Lilith (hypothetical moon)
Notes[edit]
Jump up ^ On July 8, 2013, French astrophotographer Thierry Legault successfully photographed the new Moon, although the crescent itself was invisible to the unaided eye. (astrophoto.fr)
Jump up ^ Annual aberration is the ratio of Earth's orbital velocity (around 30 km/s) to the speed of light (about 300,000 km/s), which shifts the Sun's apparent position relative to the celestial sphere toward the west by about 1/10,000 radian. Light-time correction for the Moon is the distance it moves during the time it takes its light to reach Earth divided by the Earth-Moon distance, yielding an angle in radians by which its apparent position lags behind its computed geometric position. Light-time correction for the Sun is negligible because it is almost motionless relative to the barycenter (center-of-mass) of the solar system during the 8.3 minutes that light travels between Sun and Earth. The aberration of light for the Moon is also negligible (the center of the Earth moves too slowly around the Earth-Moon barycenter (0.002 km/s); and the so-called diurnal aberration, caused by the motion of an observer on the surface of the rotating Earth (0.5 km/s at the equator) can be neglected. Although aberration and light-time are often combined as planetary aberration, Meeus separated them (op.cit. p.210).
Jump up ^ Apparent mean solar longitude is −20.496" from mean geometric longitude; apparent mean lunar longitude −0.704" from mean geometric longitude; correction to D = Moon − Sun is −0.704" + 20.496" = +19.792" that the apparent Moon is ahead of the apparent Sun; divided by 360×3600"/circle is 1.527×10−5 part of a circle; multiplied by 29.53... days for the Moon to travel a full circle with respect to the Sun is 0.000451 days that the apparent Moon reaches the apparent Sun ahead of time.
Jump up ^ see e.g. "Archived copy". Archived from the original on 2007-02-02. Retrieved 2006-12-17. CS1 maint: Archived copy as title (link) ; the IERS is the official source for these numbers; they provide TAI−UTC here and UT1−UTC here; ΔT = 32.184s + (TAI−UTC) − (UT1−UTC)
Jump up ^ delay is − (−5.8681") / (60×60×360 "/circle) / (36525/29.530... lunations per Julian century)2 × (29.530... days/lunation) days
Jump up ^ −5.8681" + 0.5×(−25.858 − −23.8946)
Jump up ^ 31 s / (86400 s/d) / [(36525 d/cy) / (29.530... d/lunation)]2
Jump up ^ from differences of various earlier determinations of the tidal acceleration, see e.g. Stephenson 1997 op.cit. par.2.2.3
References[edit]
Jump up ^ Meeus, Jean (1991). Astronomical Algorithms. Willmann-Bell. ISBN 0-943396-35-2.
Jump up ^ "new moon". Oxford English Dictionary (3rd ed.). Oxford University Press. September 2005. (Subscription or UK public library membership required.)
Jump up ^ Espenak, Fred. "Eclipses and the Moon's Orbit". NASA Eclipse Web Site. NASA. Retrieved 11 December 2016.
Jump up ^ Jawad, Ala'a H. (November 1993). Roger W. Sinnott, ed. "How Long Is a Lunar Month?". Sky&Telescope: 76..77.
Jump up ^ Meeus, Jean (2002). The duration of the lunation, in More Mathematical Astronomy Morsels. Willmann-Bell, Richmond VA USA. pp. 19..31. ISBN 0-943396-74-3.
Jump up ^ formula 47.1 in Jean Meeus (1991): Astronomical Algorithms (1st ed.) ISBN 0-943396-35-2
Jump up ^ M.Chapront-Touzé, J. Chapront (1988): "ELP2000-85: a semianalytical lunar ephemeris adequate for historical times". Astronomy & Astrophysics 190, 342..352
Jump up ^ J.Chapront, M.Chapront-Touzé, G. Francou (2002): "A new determination of lunar orbital parameters, precession constant, and tidal acceleration from LLR measurements". Astronomy & Astrophysics 387, 700–709
Jump up ^ Derived Constant No. 14 from the IAU (1976) System of Astronomical Constants (proceedings of IAU Sixteenth General Assembly (1976): Transactions of the IAU XVIB p.58 (1977)); or any astronomical almanac; or e.g. Astronomical units and constants
Jump up ^ formula in: G.M.Clemence, J.G.Porter, D.H.Sadler (1952): "Aberration in the lunar ephemeris", Astronomical Journal 57(5) (#1198) pp.46..47; but computed with the conventional value of 384400 km for the mean distance which gives a different rounding in the last digit.
Jump up ^ F.R. Stephenson, Historical Eclipses and Earth's Rotation. Cambridge University Press 1997. ISBN 0-521-46194-4 . p.507, eq.14.3
Jump up ^ Stephenson 1997 op.cit. p.38 eq.2.8
Jump up ^ Stephenson 1997 op.cit. par.14.8
Jump up ^ Lunation number in ScienceWorld
Jump up ^ "new moon". Oxford English Dictionary (3rd ed.). Oxford University Press. September 2005. (Subscription or UK public library membership required.)
Jump up ^ "Senate body decides to draft Ruet-e-Hilal Constitution".
Jump up ^ Fiqh Council of North America Decision: "Astronomical Calculations and Ramadan"
Jump up ^ Islamic Society of North America Decision:"Revised ISNA Ramadan and Eid Announcement"
Jump up ^ "A Buddhist Perspective on Fasting". www.urbandharma.org. Retrieved 2016-09-18.
Jump up ^ Posner, Menachem (2016). "How Does the Spring Equinox Relate to the Timing of Passover?". Chabad. Retrieved 9 December 2016.
Jump up ^ Momen, Moojan (2014). The Badí` (Bahá'í) Calendar: An Introduction.
Jump up ^ The Universal House of Justice (2014-07-10). "To the Bahá'ís of the World". Retrieved 2015-01-01.
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text:
RNA editing
From Wikipedia, the free encyclopedia
Jump to: navigation, search
RNA editing is a molecular process through which some cells can make discrete changes to specific nucleotide sequences within a RNA molecule after it has been generated by RNA polymerase. RNA editing is relatively rare, and common forms of RNA processing (e.g. splicing, 5'-capping, and 3'-polyadenylation) are not usually included as editing. Editing events may include the insertion, deletion, and base substitution of nucleotides within the edited RNA molecule.
RNA editing has been observed in some tRNA, rRNA, mRNA, or miRNA molecules of eukaryotes and their viruses, archaea, and prokaryotes.[1] RNA editing occurs in the cell nucleus and cytosol, as well as within mitochondria and plastids. In vertebrates, editing is rare and usually consists of a small number of changes to the sequence of affected molecules. In other organisms, extensive editing (pan-editing) can occur; in some cases the majority of nucleotides in a mRNA sequence may result from editing.
RNA-editing processes show great molecular diversity, and some appear to be evolutionarily recent acquisitions that arose independently. The diversity of RNA editing phenomena includes nucleobase modifications such as cytidine (C) to uridine (U) and adenosine (A) to inosine (I) deaminations, as well as non-templated nucleotide additions and insertions. RNA editing in mRNAs effectively alters the amino acid sequence of the encoded protein so that it differs from that predicted by the genomic DNA sequence.[2]
The Editosome Complex
Contents
[hide]
1 Editing by insertion or deletion
2 Editing by deamination
2.1 C-to-U editing
2.2 A-to-I editing
2.3 Alternative mRNA editing
3 RNA editing in plant mitochondria and plastids
4 RNA editing in viruses
5 Origin and evolution of RNA editing
6 RNA editing may be involved in RNA degradation
7 References
8 External links
Editing by insertion or deletion[edit]
The effect of uracil insertion in pre-mRNA transcripts
RNA editing through the addition and deletion of uracil has been found in kinetoplasts from the mitochondria of Trypanosoma brucei[3] Because this may involve a large fraction of the sites in a gene, it is sometimes called "pan-editing" to distinguish it from topical editing of one or a few sites.
Pan-editing starts with the base-pairing of the unedited primary transcript with a guide RNA (gRNA), which contains complementary sequences to the regions around the insertion/deletion points. The newly formed double-stranded region is then enveloped by an editosome, a large multi-protein complex that catalyzes the editing.[4][5] The editosome opens the transcript at the first mismatched nucleotide and starts inserting uridines. The inserted uridines will base-pair with the guide RNA, and insertion will continue as long as A or G is present in the guide RNA and will stop when a C or U is encountered.[6][7] The inserted nucleotides cause a frameshift, and result in a translated protein that differs from its gene.
The mechanism of the editosome involves an endonucleolytic cut at the mismatch point between the guide RNA and the unedited transcript. The next step is catalyzed by one of the enzymes in the complex, a terminal U-transferase, which adds Us from UTP at the 3' end of the mRNA.[8] The opened ends are held in place by other proteins in the complex. Another enzyme, a U-specific exoribonuclease, removes the unpaired Us. After editing has made mRNA complementary to gRNA, an RNA ligase rejoins the ends of the edited mRNA transcript.[9][10] As a consequence, the editosome can edit only in a 3' to 5' direction along the primary RNA transcript. The complex can act on only a single guide RNA at a time. Therefore, a RNA transcript requiring extensive editing will need more than one guide RNA and editosome complex.
Editing by deamination[edit]
C-to-U editing[edit]
The effect of C-to-U RNA editing on the human ApoB gene
The editing involves cytidine deaminase that deaminates a cytidine base into a uridine base. An example of C-to-U editing is with the apolipoprotein B gene in humans. Apo B100 is expressed in the liver and apo B48 is expressed in the intestines. In the intestines, the mRNA has a CAA sequence edited to be UAA, a stop codon, thus producing the shorter B48 form. C-to-U editing often occurs in the mitochondrial RNA of flowering plants. Different plants have different degrees of C-to-U editing; eight editing events occur in mitochondria of the moss Funaria hygrometrica, whereas over 1,700 editing events occur in the lycophytes Isoetes engelmanii.[11] C-to-U editing is performed by members of the pentatricopeptide repeat (PPR) protein family. Angiosperms have large PPR families, acting as trans -factors for cis -elements lacking a consensus sequence; Arabidopsis has around 450 members in its PPR family. There have been a number of discoveries of PPR proteins in both plastids and mitochondria.[12]
A-to-I editing[edit]
A-to-I editing is the main form of RNA editing in mammals[13] and occurs in regions of double-stranded RNA (dsRNA). Adenosine deaminases acting on RNA (ADARs) are the RNA-editing enzymes involved in the hydrolytic deamination of Adenosine to Inosine (A-to-I editing). A-to-I editing can be specific (a single adenosine is edited within the stretch of dsRNA) or promiscuous (up to 50% of the adenosines are edited). Specific editing occurs within short duplexes (e.g., those formed in an mRNA where intronic sequence base pairs with a complementary exonic sequence), while promiscuous editing occurs within longer regions of duplex (e.g., pre- or pri-miRNAs, duplexes arising from transgene or viral expression, duplexes arising from paired repetitive elements). There are many effects of A-to-I editing, arising from the fact that it behaves as if it is G both in translation and when forming secondary structures. These effects include alteration of coding capacity,[14] altered miRNA or siRNA target populations, heterochromatin formation, nuclear sequestration, cytoplasmic sequestration, endonucleolytic cleavage by Tudor-SN, inhibition of miRNA and siRNA processing, and altered splicing.
Alternative mRNA editing[edit]
Alternative U-to-C mRNA editing was first reported in WT1 (Wilms Tumor-1) transcripts,[15] and non-classic G-A mRNA changes were first observed in HNRNPK (heterogeneous nuclear ribonucleoprotein K) transcripts in both malignant and normal colorectal samples.[16] The latter changes were also later seen alongside non-classic U-to-C alterations in brain cell TPH2 (tryptophan hydroxylase 2) transcripts.[17] Although the reverse amination might be the simplest explanation for U-to-C changes, transamination and transglycosylation mechanisms have been proposed for plant U-to-C editing events in mitochondrial transcripts.[18] A recent study reported novel G-to-A mRNA changes in WT1 transcripts at two hotspots, proposing the APOBEC3A (apolipoprotein B mRNA editing enzyme, catalytic polypeptide 3A) as the enzyme implicated in this class of alternative mRNA editing.[19] It was also shown that alternative mRNA changes were associated with canonical WT1 splicing variants, indicating their functional significance.
RNA editing in plant mitochondria and plastids[edit]
It has been shown in previous studies that the only types of RNA editing seen in the plants' mitochondria and plastids are conversion of C-to-U and U-to-C (very rare).[20][21][22][23][24][25][26][27][28][29][30][31][32] RNA-editing sites are found mainly in the coding regions of mRNA, introns, and other non-translated regions.[22] In fact, RNA editing can restore the functionality of tRNA molecules.[24][25] The editing sites are found primarily upstream of mitochondrial or plastid RNAs. While the specific positions for C to U RNA editing events have been fairly well studied in both the mitochondrion and plastid,[33] the identity and organization of all proteins comprising the editosome have yet to be established. Members of the expansive PPR protein family have been shown to function as trans-acting factors for RNA sequence recognition.[34] Specific members of the MORF (Multiple Organellar RNA editing Factor) family are also required for proper editing at several sites. As some of these MORF proteins have been shown to interact with members of the PPR family, it is possible MORF proteins are components of the editosome complex.[35] An enzyme responsible for the trans- or deamination of the RNA transcript remains elusive, though it has been proposed that the PPR proteins may serve this function as well.
RNA editing is essential for the normal functioning of the plant's translation and respiration activity. Editing can restore the essential base-pairing sequences of tRNAs, restoring functionality.[36] It has also been linked to the production of RNA-edited proteins that are incorporated into the polypeptide complexes of the respiration pathway. Therefore, it is highly probable that polypeptides synthesized from unedited RNAs would not function properly and hinder the activity of both mitochondria and plastids.
C-to-U RNA editing can create start and stop codons, but it cannot destroy existing start and stop codons. A cryptic start codon is created when the codon ACG is edited to AUG.
Summary of the Various Functions of RNA Editing
RNA editing in viruses[edit]
RNA editing in viruses (i.e., measles, mumps, or parainfluenza) is used for stability and generation of protein variants.[37][38] Viral RNAs are transcribed by a virus-encoded RNA-dependent RNA polymerase, which is prone to pausing and "stuttering" at certain nucleotide combinations. In addition, up to several hundred non-templated A's are added by the polymerase at the 3' end of nascent mRNA.[39] These As help stabilize the mRNA. Furthermore, the pausing and stuttering of the RNA polymerase allows the incorporation of one or two Gs or As upstream of the translational codon.[39] The addition of the non-templated nucleotides shifts the reading frame, which generates a different protein.
Origin and evolution of RNA editing[edit]
The RNA-editing system seen in the animal may have evolved from mononucleotide deaminases, which have led to larger gene families that include the apobec-1 and adar genes. These genes share close identity with the bacterial deaminases involved in nucleotide metabolism. The adenosine deaminase of E. coli cannot deaminate a nucleoside in the RNA; the enzyme's reaction pocket is too small to for the RNA strand to bind to. However, this active site is widened by amino acid changes in the corresponding human analog genes, APOBEC1 and ADAR, allowing deamination.[40][41] The gRNA-mediated pan-editing in trypanosome mitochondria, involving templated insertion of U residues, is an entirely different biochemical reaction. The enzymes involved have been shown in other studies to be recruited and adapted from different sources.[4][42] But the specificity of nucleotide insertion via the interaction between the gRNA and mRNA is similar to the tRNA editing processes in the animal and Acanthamoeba mithochondria.[43] Eukaryotic ribose methylation of rRNAs by guide RNA molecules is a similar form of modification.[44]
Thus, RNA editing evolved more than once. Several adaptive rationales for editing have been suggested.[45][46] Editing is often described as a mechanism of correction or repair to compensate for defects in gene sequences. However, in the case of gRNA-mediated editing, this explanation does not seem possible because if a defect happens first, there is no way to generate an error-free gRNA-encoding region, which presumably arises by duplication of the original gene region. This thinking leads to an evolutionary proposal called "constructive neutral evolution" in which the order of steps is reversed, with the gratuitous capacity for editing preceding the "defect".[47] 31
RNA editing may be involved in RNA degradation[edit]
A study looked at the involvement of RNA editing in RNA degradation.[48] The researchers specifically looked at the interaction between ADAR and UPF1, an enzyme involved in the nonsense-mediated mRNA decay pathway (NMD). They found that ADAR and UPF1 are found within the suprasliceosome and they form a complex that leads to the down-regulation of specific genes. The exact mechanism or the exact pathways that these two are involved in are unknown at this time. The only fact that this research has shown is that they form a complex and down-regulate specific genes.
References[edit]
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Jump up ^ Malek, O., La«ttig, K., Hiesel, R., Brennicke, A. and Knoop, V. (1996). "RNA editing in bryophytes and a molecular phylogeny of land plants". EMBO J. 15 (6): 1403–1411. PMC 450045  . PMID 8635473. CS1 maint: Multiple names: authors list (link)
Jump up ^ Freyer, R., Kiefer-Meyer, M.C., and Kössel, H. (1997). "Occurrence of plastid RNA editing in all major lineages of land plants". Proc. Natl. Acad. Sci. USA. 94 (12): 6285–6296. PMC 21041  . PMID 9177209. doi:10.1073/pnas.94.12.6285. CS1 maint: Multiple names: authors list (link)
Jump up ^ Dietrich, A., Small, I., Cosset, A., Weil, J.H., and Marechal- Drouard, L. (1996). "Editing and import: Strategies for providing plant mitochondria with a complete set of functional transfer RNAs". Biochimie. 78: 518–529. doi:10.1016/0300-9084(96)84758-4. CS1 maint: Multiple names: authors list (link)
Jump up ^ Bock, R., Hermann, M., and Fuchs, M. (1997). "Identification of critical nucleotide positions for plastid RNA editing site recognition". RNA. 3: 1194–1299. CS1 maint: Multiple names: authors list (link)
Jump up ^ Gray, M.W. & Covello, P.S. (1993). "RNA editing in plant mitochondria and chloroplasts". FASEB J. 7 (1): 64–71. PMID 8422976.
Jump up ^ Marchfelder, A., Binder, S., Brennicke, A., and Knoop, V. (1998). "Preface". In: Modifcation and Editing of RNA (Grosjean, H. and Benne, R., Eds.). ASM Press, Washington, DC: 307–323. CS1 maint: Multiple names: authors list (link)
Jump up ^ Takenaka M, Zehrmann A, Verbitskiy D, Hartel B, Brennicke A (2013). "RNA Editing in Plants and Its Evolution". Annual Review of Plant Biology. 47: 335–352. doi:10.1146/annurev-genet-111212-133519.
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Jump up ^ Price, D.H. & Gray, M.W. (1998). "Editing of tRNA". In: Modification and Editing of RNA (Grosjean, H. and Benne, R., Eds.). ASM Press, Washington, DC: 289–306.
Jump up ^ Curran, J., Boeck, R., and Kolakofsky, D. (1991). "The Sendai virus P gene expresses both an essential protein and an inhibitor of RNA synthesis by shuffling modules via mRNA editing". EMBO J. 10 (10): 3079–3085. PMC 453024  . PMID 1655410. CS1 maint: Multiple names: authors list (link)
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^ Jump up to: a b Kolakofsky, D. & Hausmann, S. (1998). "Cotranscriptional paramyxovirus mRNA editing: a contradiction in terms?". In: Modification and Editing of RNA (Grosjean, H. and Benne, R., Eds.). ASM Press, Washington, DC: 413–420.
Jump up ^ Carter, C.W. (1998). "Nucleoside deaminases for cytidine and adenosine: comparisons with deaminases acting on RNA". In: Modification and Editing of RNA (Grosjean, H. and Benne, R., Eds.). ASM Press, Washington, DC: 363–376.
Jump up ^ Navaratnam, N., Fujino, T., Bayliss, J., Jarmuz, A., How, A., Richardson, N., Somasekaram, A., Bhattacharya, S., Carter, C. and Scott, J. (1998). "Escherichia coli cytidine deaminase provides a molecular model for ApoB RNA editing and a mechanism for RNA substrate recognition". J. Mol. Biol. 275 (4): 695–714. PMID 9466941. doi:10.1006/jmbi.1997.1506. CS1 maint: Multiple names: authors list (link)
Jump up ^ Covello, P.S. & Gray, M.W. (1993). "On the evolution of RNA editing". Trends Genet. 9 (8): 265–268. PMID 8379005. doi:10.1016/0168-9525(93)90011-6.
Jump up ^ Lonergan, K.M. & Gray, M.W. (1993). "Predicted editing of additional transfer RNAs in Acanthamoeba castellanii mitochondria". Nucleic Acids Res. 21 (18): 4402. PMC 310088  . PMID 8415006. doi:10.1093/nar/21.18.4402.
Jump up ^ Bachellerie, J.-P. & Cavaille, J. (1998). "Small nucleolar RNAs guide the ribose methylations of eukaryotic rRNAs". In: Modification and Editing of RNA (Grosjean, H. and Benne, R., Eds.). ASM Press, Washington, DC: 255–272.
Jump up ^ Speijer, D. (March 7, 2011). "Does constructive neutral evolution play an important role in the origin of cellular complexity?: Making sense of the origins and uses of biological complexity". BioEssays. 33 (5): 344–9. PMID 21381061. doi:10.1002/bies.201100010.
Jump up ^ Witzany G (2011). "The agents of natural genome editing". Journal of Molecular Cell Biology. 3: 181–189. PMID 21459884. doi:10.1093/jmcb/mjr005.
Jump up ^ Stoltzfus, A. (1999). "On the possibility of constructive neutral evolution". J Mol Evol. 49 (2): 169–181. PMID 10441669. doi:10.1007/PL00006540.
Jump up ^ Agranat, L., Raitskin, O., Sperling, J., and Sperling, R. (2008). "The Editing Enzyme ADAR1 and The mRNA Surveillance Protein hUpf1 Interact in the Cell Nucleus". Proc. Natl. Acad. Sci. USA. 105 (13): 5028–5033. PMC 2278206  . PMID 18362360. doi:10.1073/pnas.0710576105. CS1 maint: Multiple names: authors list (link)
External links[edit]
RNA editing website
DARNED (DAtabase of RNa EDiting in humans)
A-to-I editing website
C-to-U Editing Website
[hide]
v
t
e
Post-transcriptional modification
Nuclear
Precursor mRNA
5′ cap formation
Polyadenylation
CPSF
CstF
PAP
PAB2
CFI
CFII
Poly(A)-binding protein
RNA editing
Polyuridylation
RNA splicing
Intron / Exon
snRNP
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minor
U1
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pre-mRNA factors
PLRG1
PRPF3
PRPF4
PRPF4B
PRPF6
PRPF8
PRPF18
PRPF19
PRPF31
PRPF38A
PRPF38B
PRPF39
PRPF40A
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DCP1A
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