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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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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 5/04/2023 07:24:00 AM, Blogger Hemmaharry said...

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