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June 16, 2006

Fruit Genetics Friday #3: Self-Incompatibility

When shopping for fruit and nut trees (or just flipping idly through nursery catalogs), one common issue is that many species require another of their species to provide pollen in order to set fruit. Most fruit crops produce flowers which are both male and female, and produce plenty of pollen. In some cases odd ploidy can result in plants incapable of producing viable pollen at all (or very small amounts, anyway), and a few species are dioecious (having male and female flowers on separate plants). But in many cases, the plants in question produce perfectly good pollen. Nursery catalogs are full of little pollination tables and warnings of "requires a pollinator" or, strangely enough, something along the lines of "[Cultivar X] will not pollinate [Cultivar Y]".

The reason for this is a phenomenon called self-incompatibility (SI). As inconvenient as it is for those of us who would like to just plant our favorite cultivar and nothing else, self-incompatibility serves a useful purpose evolutionarily. The whole point of sexual reproduction is to encourage assortment and exchange of genes, and yet by generally self-fertilizing (which they do by having male and female reproductive organs mature simultaneously and adjacent to each other), many species have sacrificed this evolutionary opportunity in exchange for a better chance of producing offspring, with the downside that those offspring will be merely reassortments of the parent's genes, not new mixtures of two individuals. Self-incompatibility evolved to prevent self-pollination in such cases, helping to avoid inbreeding depression.

There are two major SI systems in fruit crops, and both are really quite simple, though the implications are not necessarily so. Both involve interactions involve alleles of a single gene, called the "S" locus, which is why the basic genetics of it were worked out nearly a hundred years ago.

Gametophytic self-incompatibility, the most common form, is most commonly seen in sweet cherry (Prunus avium, which, coincidentally, I'm eating a giant bag of right now), though it's also found in a number of other fruit and nut species including almond, Japanese apricot, roses, arctic bramble and assorted diploid blackberries and raspberries. Under this scheme, the haploid genotype of the pollen is the factor determining whether pollination can occur. Each pollen grain receives one of the two S-alleles of the parent, and that pollen is unable to germinate on flowers of individuals which possess the same S-allele. So, for example:

If you have a cherry tree with the S-genotype of S1S2, the pollen grains produced will be either S1 or S2. Neither can fertilize the tree's own flowers, since both S1 and S2 are present in the female. But say a neighoring tree is S1S3. It can produce S1 and S3 pollen. The S1 grains won't be able to germinate on either tree's flowers, but the S2 grains of the first tree can fertilize the second, and the second's S3 grains can fertilize the first. If we propose third tree, S4S8, appears on the scene, then both the other plants, having neither S4 or S8, will be able to fertilize it, as well as the reverse. There are quite a few different S-alleles in cherry, so the odds of getting two cultivars with the exact same complement is pretty small, but it can happen (for example, 'Bing' and 'Emperor Francis' are both S3S4). A nice diagram of gametophytic SI is available here.

The other system is called sporophytic incompatibility. This one's a bit less common, particularly in fruit. The example that springs to mind is hazelnut (Corylus species). The situation is similar to the gametophytic system, in that it depends on a pair of S-alleles. But there is one critical difference: in gametophytic SI, the compatibility phenotype of the pollen grain depends on the genotype of that grain, while under sporophytic it is the genotype of the parent that matters. So while a S1S2 GSI plant produces pollen grains with S1 or S2 genotypes, and corresponding S1 or S2 compatibility phenotypes. However, under SSI, while pollen grains are still genotypically S1 or S2, they all will share the dominant S-alleles phenotype (in this case S1). So a S1S2 SSI plant would produce pollen grains carrying the S2 allele, but effectively S1 for pollination purposes, so it could pollinate a S2S3 plant. An interesting twist is that while GSI plants are always heterozygous (with one minor exception which I will get to), SSI plants can carry two identical S-alleles, because it the actual genotype of the pollen is not related to the phenotype. It also adds the extra complication of the needing to know the dominance hierarchy of the S-alleles (which, inconveniently, is not simply numerical order. It's actually something like: (3, 8), (6), (1, 5, 7, 10, 12, 14, 15, 16, 17, 18, 20, 21, 24), (2, 25), (19), (9, 11, 22, 26), (23), (4), with dominance running from dominant on the left to recessive on the right (alleles in the same set of parentheses are roughly equivalent in dominance).

Researchers have, however, managed to come up with a convenient end-run around GSI in cherry. Scientists pollinated 'Emperor Francis' (S3S4) with irradiated pollen from 'Napoleon' (also S3S4). Normally, with both parents having the same S-alleles, no cross could occur. But a few seeds did set, because the radiation had damaged the chromosomes in some of the pollen grains, and, completely randomly, had damaged the S4 allele carried in one of them. The resulting seedling was JI 2420, which broke the rule I just mentioned above by being S4S4. However, one of those S4's was non-functional, and could pollinate any sweet cherry, including itself. The value of this was obvious, and JI 2420's amazing allele was immediately incorporated into a new cultivar, called 'Stella'. Nearly all self-compatible sweet cherries carry S4', the modified S4 from 'Stella'. While the trait is undeniably useful, the dependence on a single source is worrisome from a genetic diversity standpoint, and Amy Iezzoni, cherry breeder from Michigan State, has suggested that there may be some undesireable linkages associated with Stella's S4' gene. (Another self-compatible selection, JI 2434, came from the same experiment as JI 2420 and has been found to carry a second self-compatibility gene, this one a modification of the S3 allele (cleverly indicated S3'). So there's at least another option out there. Similar self-compatibility alleles exist in almond, Japanese apricot, and probably Rubus, having occurred naturally and been selected.

So next time you see those compatibility tables in the nursery catalog, see if you can puzzle out the S-allele genotypes based on the information given. And don't forget to buy a pollenizer!

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At 3/21/2012 03:35:00 PM, Blogger den_den said...

Funny Storycomo ganhar dinheiro na internetFraming all this was his career as a writer and editor. After moving to England, Pevsner had found that the study of architectural history had little status in academic circles, and the amount of information available, especially to travellers wanting to inform themselves about the architecture of a particular district, was limited. Invited by Allen Lane, founder of Penguin Books, for whom he had written his Outline and also edited the King Penguin series,[8] to suggest ideas for future publications, he proposed a series of comprehensive county guides to rectify this shortcoming.

At 3/26/2012 09:03:00 PM, Blogger camap said...

The depression tracked westward as it intensified into a tropical storm, and after passing north Puerto Rico, Eloise briefly attained hurricane intensity.Denver Personal Injury Attorney box trailers brisbane

At 3/26/2012 09:20:00 PM, Blogger camap said...

. As Eloise progressed westward, it affected Cuba, though to a lesser extent. Upon making landfall in Florida, Eloise generated winds reportedly gusting to 155 mph (249 km/h). cubicles Decatur chiropractor

At 3/26/2012 09:43:00 PM, Blogger camap said...

. Breeding males are brightly coloured with red or orange and black plumage. The non-breeding male and female have streaky brown plumage, paler below.regenboog carwrapfolie Cardiff minibus

At 3/26/2012 11:45:00 PM, Blogger camap said...

The depression transitioned into an extratropical storm over Virginia, and became indistinguishable by later that same day. The remnant moisture, however, merged with a weather front to produce widespread and heavy precipitation.[1][2]OHSAS 18001 engineering companies brisbane

At 3/30/2012 05:42:00 AM, Blogger camap said...

In advance of Hurricane Eloise, warnings for heavy rainfall and potential flooding were issued for Puerto Rico and the Virgin Islands. A hurricane warning was declared for parts of the Dominican Republic about 12 hours before landfall.[hens night brisbane breitling replica

At 3/30/2012 07:17:00 PM, Blogger amantr said...

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At 4/07/2012 03:24:00 PM, Blogger yeu_yeu said...

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Flash cartoonsDaniel Boone (November 2, 1734 [O.S. October 22] – September 26, 1820) was an American pioneer, explorer, and frontiersman whose frontier exploits made him one of the first folk heroes of the United States.

At 4/07/2012 04:32:00 PM, Blogger yeu_yeu said...

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estojo original rolexMany parts of the Americas are still populated by indigenous Americans; some countries have sizable populations, especially Bolivia, Peru, Mexico, Guatemala, Colombia, and Ecuador. At least a thousand different indigenous languages are spoken in the Americas.[33] Some, such as Quechua languages, Aymara, Guaraní, Mayan languages, and Nahuatl, count their speakers in millions.

At 4/13/2012 07:46:00 PM, Blogger aloalo said...

The highest known mountain in the Solar System is Olympus Mons on the planet Mars at 21,171 m (69,459 ft). Mountains and mountain ranges on Earth are typically formed by the movement and/or interaction of lithospheric plates.
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At 4/16/2012 05:19:00 PM, Blogger yeu_yeu said...

Brazil's current foreign policy is based on the country's position as a regional power in Latin America, a leader among developing countries, and an emerging world power.[183] In general, current Brazilian foreign policy reflects multilateralism, peaceful dispute settlement, and nonintervention in the affairs of other countries.
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At 4/18/2012 07:17:00 PM, Blogger yeu_yeu said...

NATO defines Indirect fire as "Fire delivered at a target which cannot be seen by the aimer."[1] The implication is that azimuth and/or elevation ‘aiming’ is done using instrumental methods. Hence indirect fire means applying ‘firing data’ to azimuth and elevation sights and laying these sights. Indirect fire uses a high trajectory, which further distinguishes it from direct fire.
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At 4/22/2012 04:21:00 PM, Blogger yeu_yeu said...

In the early years of the 20th century (1907 to 1914) the Deutsche Orientgesellschaft expedition, led by Ludwig Borchardt, excavated extensively throughout the North and South suburbs of the city. The famous bust of Nefertiti, now in Berlin's Ägyptisches Museum, was discovered amongst other sculptural ar
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At 4/28/2012 01:22:00 AM, Blogger aloalo said...

German battleship construction. Admiral John Fisher, who became the First Sea Lord and head of the Admiralty in 1904, introduced sweeping reforms in large part to counter the growing threat posed by the expanding German fleet. Training programs were modernized, old and obsolete vessels were discarded, and the scattered squadrons of battleships were consolidated into four main fleets, three of which were based in Europe. Britain also made a series of diplomatic arrangements, including an alliance with Japan that allowed a greater concentration of British battleships in the North Sea.
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At 4/28/2012 04:11:00 AM, Blogger aloalo said...

Scheer ordered the fleet on sweeps of the North Sea on 26 March, 2–3 April, and 21–22 April. The battlecruisers conducted another raid on the English coast on 24–25 April, during which the fleet provided distant support.[42] Scheer planned another raid for mid-May, but the battlecruiser Seydlitz had struck a mine during the previous raid and the repair work forced the operation to be pushed back until the end of the month.
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At 4/28/2012 09:49:00 AM, Blogger HT said...

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I must confess to being a bit rubbish at gadgets - my current mobile is the sort that people snigger at openly as it is *sssshhh* quite old and not very cool and can't take photos or anything fancy like that.

At 4/28/2012 10:45:00 AM, Blogger HT said...

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must confess to being a bit rubbish at gadgets - my current mobile is the sort that people snigger at openly as it is *sssshhh* quite old and not very cool and can't take photos or anything fancy like that.

At 4/28/2012 08:04:00 PM, Blogger trung said...

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At 4/28/2012 10:42:00 PM, Blogger LU-PICACHU said...


What is Capsiplex? must confess to being a bit rubbish at gadgets - my current mobile is the sort that people snigger at openly as it is *sssshhh* quite old and not very cool and can't take photos or anything fancy like that.

At 4/29/2012 01:15:00 AM, Blogger trung said...

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it isn't hard to find.
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It always seems to be so hard to give.

At 4/30/2012 06:03:00 PM, Blogger aloalo said...

After another billion years all surface water will have disappeared[26] and the mean global temperature will reach 70 °C[60] (158 °F). The Earth is expected to be effectively habitable for about another 500 million years from that point,[25] although this may be extended up to 2.3 billion years if the nitrogen is removed from the atmosphere.[27] Even if the Sun were eternal and stable, the continued internal cooling of the Earth would result in a loss of much of its CO2 due to reduced volcanism,[61] and 35% of the water in the oceans would descend to the mantle due to reduced steam venting from mid-ocean ridges.
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At 4/30/2012 06:57:00 PM, Blogger aloalo said...

Earth's biosphere has significantly altered its atmosphere. Oxygenic photosynthesis evolved 2.7 billion years ago, forming the primarily nitrogen-oxygen atmosphere of today. This change enabled the proliferation of aerobic organisms as well as the formation of the ozone layer which blocks ultraviolet solar radiation, permitting life on land.
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At 5/01/2012 12:51:00 AM, Blogger aloalo said...

The band made their public debut in a one-off performance in April 1973, and featured Robert Smith (piano), Michael "Mick" Dempsey (guitar), Laurence "Lol" Tolhurst (percussion), Marc Ceccagno (lead guitar) and Alan Hill (bass guitar).[6] In January 1976 the band took a more substantial form of what would be when Ceccagno formed Malice with Smith and Dempsey along with two other classmates from St. Wilfrid's Catholic Comprehensive School, with Ceccagno on lead, Smith on guitar, and Dempsey switching to bass. Ceccagno soon left, however, to form a jazz-rock fusion band called Amulet.
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At 5/01/2012 06:06:00 AM, Blogger aloalo said...

The WTO's predecessor, the General Agreement on Tariffs and Trade (GATT), was established after World War II in the wake of other new multilateral institutions dedicated to international economic cooperation — notably the Bretton Woods institutions known as the World Bank and the International Monetary Fund. A comparable international institution for trade, named the International Trade Organization was successfully negotiated.
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At 5/02/2012 05:45:00 AM, Blogger aloalo said...

Pharmacopoeia (written in Latin, and used for regulating the composition of medicines from 1618 and, through the College's police the Censors, for enforcing the College's monopoly on medical science, then being challenged by the Society of Apothecaries), and the `Nomenclature of Diseases' in 1869. The latter created the international standard for the classification of diseases which was to last until the World Health Organisation's Manual of the international classification of diseases superseded it in the twentieth century.
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At 5/03/2012 05:16:00 AM, Blogger aloalo said...

Following concerns that the South Island might form a separate colony, premier Alfred Domett moved a resolution to transfer the capital from Auckland to a locality near the Cook Strait.[46] Wellington was chosen for its harbour and central location, with parliament officially sitting there for the first time in 1865. As immigrant numbers increased, conflicts over land led to the New Zealand Wars of the 1860s and 1870s, resulting in the loss and confiscation of much Māori land.
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At 5/03/2012 05:52:00 AM, Blogger aloalo said...

Because of financial pressures and the desire to consolidate railways, education, land sales and other policies, government was centralised and the provinces were abolished in 1876.[106] As a result, New Zealand now has no separately represented subnational entities. The provinces are remembered in regional public holidays[107] and sporting rivalries.
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At 5/04/2012 11:45:00 AM, Blogger aloalo said...

For a number of years a creamery was operated for farmers who brought their milk in daily to be separated. (The stone structure which once housed the creamery can be seen today.) The cream was shipped to Baltimore while the skim milk was used by the farmers on their farms. With the passage of time, Sparks' Switch came to be known simply as "Sparks." In 1888, the area had grown to a point where "a substantial foot bridge 6 feet in width" had to be built across the Gunpowder River.
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At 5/04/2012 02:55:00 PM, Blogger trung said...

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The record shows I took the blows -
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At 5/04/2012 10:35:00 PM, Blogger camap said...

arayan was born in Udaipur and learned to play the sarangi at an early age. He studied under sarangi players and singers relogios maquina ETA water ionizer

At 5/05/2012 12:51:00 AM, Blogger aloalo said...

Lahore appears as the capital of the Punjab for the first time under Anandapala – the Hindu Shahi king who is referred to as the ruler of (hakim i lahur) –after leaving the earlier capital of Waihind.[22] Few references to Lahore remain from before its capture by Sultan Mahmud of Ghazni in the eleventh century.
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At 5/05/2012 06:41:00 AM, Blogger aloalo said...

After 1876, the resulting political union was officially called the Indian Empire and issued passports under that name. As India, it was a founding member of the League of Nations, and a member nation of the Summer Olympics in 1900, 1920, 1928, 1932, and 1936.
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At 5/05/2012 08:59:00 AM, Blogger aloalo said...

reserves and the striking forces. Armed with seven-foot metal tipped lathis and smoothbore muskets, and tear gas after 1940, they repressed the disturbances of 1930-33. Special striking forces included the Malabar Special Police, armed with Enfield rifles. It was established to handle the Moplah rebellion of 1921 and was used throughout the presidency. The Presidency General Reserve was established in 1931
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At 5/05/2012 04:24:00 PM, Blogger trung said...

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At 5/06/2012 12:14:00 AM, Blogger simson11 said...

Hi Rahina!
Like everybody I am running out time not out of ideas. It is so wonderful to paint outside, although it is sometimes very exhausting to capture the fleeting moment. Thanks!
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At 5/06/2012 12:25:00 AM, Blogger M.U.P said...

Apart from its history, culture, and scenic locations, it is also known for its Rajput-era palaces. The Lake Palace, for instance, covers an entire island in the Pichola Lake. Many of the palaces have been converted into luxury hotelsdating ideas cheap insurance quotes

At 5/06/2012 12:54:00 AM, Blogger M.U.P said...

Udaipur was founded in 1559 by Maharana Udai Singh II as the final capital of the erstwhile Mewar kingdom, located to the southwest of Nagda4Life wholesale lace front wigs

At 5/06/2012 05:11:00 AM, Blogger M.U.P said...

Mornings, evenings and nights are cold with minimum temperature around 11.6 °C (52.9 °F) especially if there is a slight breeze in the air.[7]medical marijuana doctor مفارش

At 5/07/2012 12:20:00 AM, Blogger aloalo said...

The Adventures of Brisco County, Jr., about a Harvard-educated bounty hunter who wants to avenge the death of his father, the most famous lawman in the Old West. Fox gave the go ahead for the series. Brisco also had a science fiction element, in the form of a mysterious orb which appears in several episodes. Boam went back to making features, leaving Cuse to co-create and executive produce the critically acclaimed series. Afterwards, Cuse gave much of the credit for the show's success to actor Bruce Campbell who played Brisco County Jr., the lead character.
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At 5/07/2012 12:24:00 AM, Blogger aloalo said...

A team of eight top Chinese stuntmen and coordinators out of Hong Kong were hired. Stanley Tong, who had directed many of Jackie Chan's biggest Hong Kong features, directed the pilot. Sammo Hung became the first Chinese actor to star as the lead in an American TV series.[14] Cuse was running both Martial Law and Nash Bridges simultaneously. The workload became creatively and physically difficult, which lead to him leaving Martial Law. and focus exclusively on Nash Bridges. Another factor, Cuse said, were creative differences with Sammo Hung about the future direction of Martial Law.

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At 5/08/2012 04:36:00 AM, Blogger aloalo said...

John's judicial reforms had a lasting, positive impact on the English common law system, as well as providing an additional source of revenue. An argument with Pope Innocent III led to John's excommunication in 1209, a dispute finally settled by the king in 1213. John's attempt to defeat Philip in 1214 failed due to the French victory over John's allies at the battle of Bouvines. When he returned to England, John faced a rebellion by many of his barons, who were unhappy with his fiscal policies and his treatment of many of England's most powerful nobles. Although both John and the barons agreed to the Magna Carta peace treaty in 1215, neither side complied with its conditions.
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At 5/08/2012 04:42:00 AM, Blogger aloalo said...

Civil war broke out shortly afterwards, with the barons aided by Louis of France. It soon descended into a stalemate. John died of dysentery contracted whilst on campaign in eastern England during late 1216; supporters of his son Henry III went on to achieve victory over Louis and the rebel barons the following year.
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At 5/08/2012 05:45:00 AM, Blogger aloalo said...

John's first period of rule in Ireland was not a success. Ireland had only recently been conquered by Anglo-Norman forces, and tensions were still rife between Henry II, the new settlers and the existing inhabitants.[27] John infamously offended the local Irish rulers by making fun of their unfashionable long beards, failed to make allies amongst the Anglo-Norman settlers, began to lose ground militarily against the Irish and finally returned to England later in the year, blaming the viceroy, Hugh de Lacy, for the fiasco
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At 5/09/2012 05:36:00 AM, Blogger aloalo said...

Sparta is located in the region of Laconia, in the south-eastern Peloponnese. Ancient Sparta was built on the banks of the Evrotas River, the main river of Laconia, which provided it with a source of fresh water. The valley of the Evrotas is a natural fortress, bounded to the west by Mt. Taygetus (2407 m) and to the east by Mt. Parnon (1935 m). To the north, Laconia is separated from Arcadia by hilly uplands reaching 1000 m in altitude. These natural defenses worked to Sparta's advantage and contributed to Sparta never having been sacked. Though landlocked, Sparta had a harbor, Gytheio, on the Laconian Gulf.
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At 5/09/2012 11:25:00 AM, Blogger aloalo said...

Three battles called the 'Battle of Lepanto' were fought in this gulf: that of 1499, that of 1500 and the largest of the three, that of 1571. Lepanto itself is in the Gulf of Corinth, not in the Gulf of Patras. A battle of Patras was fought here in 1772 between Russian and Ottoman fleets.
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At 5/10/2012 09:54:00 AM, Blogger aloalo said...

In 1935 a backlash against the modernizing, anti-religious policies of Reza Shah erupted in the Mashed shrine. Responding to a cleric who denounced the Shah's heretical innovations, corruption and heavy consumer taxes, many bazaaris and villagers took refuge in the shrine, chanted slogans such as "The Shah is a new Yezid." For four full days local police and army refused to violate the shrine and the standoff was ended when troops from Azerbaijan arrived and broke into the shrine,[8] killing dozens and injuring hundreds, and marking a final rupture between Shi'ite clergy and the Shah.
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At 5/11/2012 12:46:00 AM, Blogger simson11 said...

Hi Rahina!
Like everybody I am running out time not out of ideas. It is so wonderful to paint outside, although it is sometimes very exhausting to capture the fleeting moment. Thanks!
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At 5/11/2012 11:04:00 PM, Blogger M.U.P said...

In middle age Hepburn found a niche playing spinsters, such as in The African Queen, and became a Shakespearean stage actress. She continued to work into old agetimeshare business for sale perth

At 5/14/2012 01:38:00 PM, Blogger aloalo said...

France in the midst of the French Revolution. In an attempt to revive their relationship, she travelled to Scandinavia on business for him, taking the one-year-old Fanny with her, but the affair never rekindled. After falling in love with and marrying the philosopher William Godwin, Wollstonecraft died in childbirth in 1797, leaving the three-year-old Fanny in the hands of Godwin, along with the newborn Mary Wollstonecraft Godwin. Four years later, Godwin remarried and his new wife,
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At 5/16/2012 12:39:00 PM, Blogger aloalo said...

A hypothesis for the phylogeography of this group has been proposed and this provides a nice example of various speciation mechanisms at work. In this scenario, ancestors of this group became isolated from the remaining parrots when New Zealand broke away from Gondwana about 82 million years ago, resulting in a physical separation of the two groups.[14][15] This mechanism is called allopatric speciation.
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At 11/25/2013 06:05:00 PM, Blogger Jay Lamptey said...

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