.comment-link {margin-left:.6em;}

Fruit Blog Header.jpg
APPLES - BRAMBLES - CITRUS - GRAPES - NUTS - PAWPAWS - PEARS - PERSIMMONS - STONE FRUITS - STRAWBERRIES - FORUM

Cherimoya
My cherimoya, shortly before it became smoothie fodder.

Archive of past images




AppleBorovinka


Chinquapin

Recent Cultivar Releases
Pomegranate

Current Work in Pomology

StrawbParkerEarle


QuinceBourgeat


Atom Feed

Powered by Blogger

Add to Technorati Favorites

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!

Labels: , , , , ,

120 Comments:

At 11/18/2011 02:51:00 AM, Anonymous Anonymous said...

Horoskopcelebrity photos
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 11/27/2011 01:25:00 AM, Anonymous Anonymous said...

payday advance loanspropane heaters
I remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review and using it as a cover quote, just to piss the reviewer in question off... I'm sure that's not the case here though!

 
At 11/30/2011 03:16:00 AM, Anonymous Anonymous said...

BEST ANDROID TABLET REVIEW
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 12/01/2011 07:17:00 AM, Anonymous Anonymous said...

building furnitureIrving Apartments
I remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review

 
At 12/04/2011 04:09:00 AM, OpenID lowderwin said...

exhibition stand manufacturerscheap motor cycle batteries
I remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review and using it as a cover quote, just to piss the reviewer in question off I'm sure that's not the case here though!

 
At 12/05/2011 03:45:00 AM, OpenID lowderwin said...

Cheap DJs in NYdesigner dog clothing
ot only Tree, but also Larry and Curtis. I think what grandpa said was right. Sometimes when your mad you want to get over with everything by letting out your anger. I think the most common thing people do is to yell out words or just hit on something hard until they feel better.

 
At 12/07/2011 01:01:00 AM, Anonymous Anonymous said...

Mercedes HIDlocation de salle marseille
In that 15-minute exchange with Fox News' Brett Baier, Romney bristled at questions about his changed views on abortion, climate change, immigration and gay rights, all of which are widely discussed in political circles.

Romney acknowledged rejecting his pro-abortion-rights stand of the 1990s, although he did not explain why. Otherwise, he told Baier, "Your list is just not accurate." Romney suggested the questions were inspired by "Democratic ads" that label him a serial flip-flopper.

 
At 1/05/2012 02:43:00 PM, Blogger mantr said...

Ignition Distributoredinburgh city hotels
And I've been keeping all the letters that I wrote to you
Each one a line or two
"I'm fine baby, how are you?"
Well I would send them but I know that it?s just not enough
My words were cold and flat
And you deserve more than that

 
At 1/11/2012 05:56:00 AM, Blogger kosovohp said...

I give good love
I'll buy your clothes
I'll cook your dinner too
Soon as I get home from work
I'll pay your rent
Your faithful lover
Audio Productioninsanity workout

 
At 2/01/2012 07:45:00 AM, Blogger CuteGirl said...

US presidential hopeful Mitt Romney has won a clear victory in the Republican primary election in Florida, official results indicate.

Mr Romney polled 46%, but rival Newt Gingrich, who only managed 32%, says he has no intention of dropping out.

Florida's primary was the biggest of the 2012 election season so far and saw a bitter TV ad battle between the men hoping to take on Barack Obama.
Auto Warrantycheap tickets to middle east

 
At 2/07/2012 01:03:00 AM, Blogger tung said...

"Cartman Gets an Anal Probe" is the pilot episode of the animated television series South Park. It first aired on Comedy Central in the United States on August 13, 1997. The episode introduces child protagonists Eric Cartman, Kyle Broflovski, Stan Marsh and Kenny McCormick, who attempt to rescue Kyle's younger brother Ike from being abducted by aliens. At the time of the writing of the episode, South Park creators Trey Parker and Matt Stone did not yet have a series contract with Comedy Central.
anger management techniqueshealth insurance florida

 
At 2/08/2012 02:15:00 PM, Blogger kosovohp said...

Must've been a silly one
To sacrifice a pot of gold
You're the kind of woman
That needs a man that's always there
It's not that you require a lot
Just need some tender love and care

HRT Womenquail west

 
At 2/10/2012 11:10:00 AM, Blogger Justlia From TGR said...

business developmentFast Computer RepairI remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review and using it as a cover quote, just to piss the

 
At 2/10/2012 11:31:00 PM, Blogger Justlia From TGR said...

holidays to maldiveshome decor ideasI must confess to being a bit rubbish at gadgets - my current mobile is the sort that people snigg

 
At 2/11/2012 12:34:00 AM, Blogger Justlia From TGR said...

cheap wedding ringsfranchise opportunitiesI remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review and using it as a cover

 
At 2/11/2012 05:26:00 AM, Blogger Justlia From TGR said...

Stacked Car InsuranceWhat is Phen375 side effects?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

 
At 2/11/2012 09:45:00 AM, Blogger Justlia From TGR said...

hair again reviewcute teen fashionI 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

 
At 2/11/2012 01:21:00 PM, Blogger Justlia From TGR said...

active directory softwarefree textingI 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 2/11/2012 01:28:00 PM, Blogger Justlia From TGR said...

free text messageactive directory recoveryI remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review

 
At 2/12/2012 05:37:00 AM, Blogger M.U.P. said...

Given the latitude, long periods of constant darkness or constant sunlight create climates unfamiliar to human beings in much of the rest of the world.Xerox 6125 toner Condos in Mississauga

 
At 2/12/2012 02:36:00 PM, Blogger Justlia From TGR said...

holidays in antiguavaporizerI 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 2/13/2012 05:50:00 PM, Blogger trungvkvk said...

build a websitewebmaster
The scars of your love remind me of us
They keep me thinking that we almost had it all
The scars of your love they leave me breathless
I can't help feeling
We could've had it all
Rolling in the deep
You had my heart inside of your hand
And you played it
To the beat

 
At 2/14/2012 05:00:00 PM, Blogger trungvkvk said...

Galapagos CruisesHotels in Shimla
Each and every time I turn around to leave
I feel my heart begin to burst and bleed
So desperately I try to link it with my head
But instead I fall back to my knees
As you tear your way right through me
I forgive you once again
Without me knowing
You've burnt my heart to stone

 
At 2/15/2012 04:18:00 AM, Blogger M.U.P. said...

members of the Upper House also controlled tiny constituencies, known as pocket or rotten boroughs, and could ensure the election of their relatives or supportersclases particulares for life

 
At 2/15/2012 05:24:00 AM, Blogger M.U.P. said...

Tummy Tuck argan yagiNo individual may be a member of both Houses, and members of the House of Lords are legally barred from voting in elections for members of the House of Commons

 
At 2/16/2012 10:35:00 AM, Blogger M.U.P. said...

All legislation must be passed by the House of Commons to become law and it controls taxation and the supply of money to the governmentgrow hair naturally abbigliamento tennis

 
At 2/16/2012 11:05:00 AM, Blogger Justlia From TGR said...

free hostingarticle writing serviceI 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 2/17/2012 04:32:00 PM, Blogger trung said...

Planktonnegle lak
I was her she was me
We were one we were free
If there's somebody calling me on
She's the one
If there's somebody calling me on
She's the one

 
At 2/18/2012 08:36:00 AM, Blogger M.U.P. said...

These bills do not become laws; they are ceremonial indications of the power of each House to debate independently of the Crown. After the pro forma bill is introduced, each House debates the content of the Speech from the Throne for several daysFlorida Drug Rehab Center custom packaging

 
At 2/19/2012 07:45:00 PM, Blogger Justlia From TGR said...

pull up banner standsDepression Treatment Los AngelesI 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 2/20/2012 02:34:00 PM, Blogger M.S said...

List of Top Car Insurance CompaniesbromfietsverzekeringI 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 2/20/2012 04:21:00 PM, Blogger trungvkvk said...

Quality Veterinary Equipmentclases particulares valencia
I heard, that you're dreams came true
I guess she gave you things
I didn't give to you

Old friend, why are you so shy
Ain't like you to hold back
Or hide from the light

 
At 2/21/2012 08:46:00 AM, Blogger carotkhonglo said...

Nowadays Paulo Coelho is one of the most read and translated authors. His bestseller The Alchemist has been translated into 73 languages and is one of the most read books in the world.load balancing Atlanta Web Design Company

 
At 2/22/2012 01:19:00 AM, Blogger carotkhonglo said...

South America relies heavily on the exporting of goods and natural resources. On an exchange rate basis Brazil (the seventh largest economy in the world and the largest in South AmericaParkett München cream

 
At 2/22/2012 03:38:00 AM, Blogger carotkhonglo said...

I kept a steady pace and felt I wasn't losing important time; according to Strava, I complete the nearly 6-mile climb, averaging 8 mph and 218 watts, well over a minute faster than last yearcomo hacer un guionElektroinstallation München

 
At 2/22/2012 09:53:00 PM, Blogger carotkhonglo said...

In 1990, Steve Jobs at Stanford University on a speech, Christian Louboutin Alta Ariella Talon Leopard Bootsnoticed the front row to listen to Lauren. That night, a meeting is hard to wait as one cause of Jobs.hypnotherapy Dorsetwebsite video production

 
At 2/23/2012 01:23:00 PM, Blogger 25251325 said...

Former Senator Rick Santorum of Pennsylvania won three Republican Party (GOP) presidential contests on Tuesday, but received no delegates. Though the races were non-binding, some analysts see in them a vulnerability for front-runner Mitt Romney, and a possible revitalization of the Santorum campaign.Capsiplex ReviewsBaltimore Limo

 
At 2/24/2012 04:36:00 AM, Blogger M.S said...

me encantan tus trabajos, son muy finos y bonitos. Y me gusta especialmentedebt collectionelectronic cigarette

 
At 2/24/2012 09:54:00 AM, Blogger carotkhonglo said...

stradbroke accommodationmarketing displaysBecause of South America's broad ethnic mix, South American cuisine has African, American Indian, Asian, and European influences. Bahia, Brazil

 
At 2/25/2012 05:48:00 AM, Blogger trungvkvk said...

iPhone App DevelopmentLinkedIn Marketing Guide
Rebel soldiers

Thousands of people are trapped in residential areas which government forces have subjected to days of heavy bombardment. The districts are defended by rebel soldiers calling themselves the Free Syrian Army, who are only lightly armed.
The injured and dying are treated in makeshift clinics with limited supplies.

 
At 2/25/2012 02:41:00 PM, Blogger trungvkvk said...

after infidelityCasas Rurales
Halfpenny missed a simple penalty chance after his pack made a mess of the English scrum, and when Dickson went on a dart after a tap-and-go, England finally made inroads

Manu Tuilagi hammered dents in the red defence and Farrell made it 3-0, only for Halfpenny to level things up after Chris Robshaw took a pass standing still and was smashed by Dan Lydiate.

 
At 2/26/2012 09:14:00 AM, Blogger picasxo said...

After twenty minutes or so, it was time to get moving, as the sun was down and the temperature was dropping fast. I wanted to get out and get moving before it got too cold. I was somewhat surprised to encounter hills after leaving the checkpoint, I thought that Minnesota was flat. I later found out that the hills would continue from thirty five miles all the way to the one hundred fifteen mile mark. I hoped to make the halfway checkpoint at Melgeorges resort in another twelve hours, but the hills and need for sleep got the better of me. There were shelters along the trail that were spaced about ten miles apart and they made good bivy spots for us. προώθηση ιστοσελίδωνprofesoresparticulares de ingles en madrid

 
At 2/26/2012 11:47:00 AM, Blogger yeu_yeu said...

As in past years, the main struggle is keeping Wayne eating. It is a constant struggle to keep up with the huge caloric expenditure. Christian, from France was the winner in the deca race, finishing in just under 20 hours, only about eight hours off of the world record. In the double deca, it is close at the top, with four men within a few hours of each other. They are still on the bike portion, with about 250 miles still to go. We are seeing lot of unusual bike problems. Gregor Sundin, from Sweden, had his rear derailluer shifter break, so we fixed the bike in a single gear so that he could continue to ride. Once he stops for a sleep break, I will try to get the shifter repaired. It may not be possible. The folks from Denmark have an entire team of Helpers, and we have borrowed a few things from them. They have a bike mechanic, but he took a few days off, so he is not around to help with repairs. They also posted some video from early in the race, you can see it by going to decatri.dk. Boston Web DesignerLearn About Personal Injury Law

 
At 2/26/2012 09:58:00 PM, Blogger carotkhonglo said...

I think in some ways we were a more open society back in the 80's. Like for instance, I remember that in Manhattan it wasn;'t at all unusal to go into an office and find the CEO and the Controller snorting coke on the Boardroom table. That was ok back then too.business it supportannonce véhicule autos

 
At 2/27/2012 06:42:00 PM, Blogger trung said...

security guards companiesExpert Credit Card Reviews
great
When the rain is blowing in your face,
and the whole world is on your case,
I could offer you a warm embrace
to make you feel my love.

 
At 2/28/2012 05:51:00 AM, Blogger tung said...

Offshoots of the imperial dynasty, some grandsons of Emperor Kammu were first given the name Taira in 825 or later. Afterwards, descendants of Emperor Nimmyō, Emperor Montoku, and Emperor Kōkō were also given the surname. The specific hereditary lines from these emperors are referred to by the emperor's posthumous name followed by Heishi. i.e. Kammu Heishi.
The Taira were one of the four important clans that dominated Japanese politics during the Heian period (794-1185) - the others were the Fujiwara, the Tachibana and the Minamoto.
charms charm braceletskeratin hair treatment

 
At 2/29/2012 05:06:00 AM, Blogger banhque10k said...

I went over to take some pictures of the asshats, and get a closer look at them. I can report that they appeared to be mono-gastric, bi-pedal, and carbon-based; other than that, I really couldn't see even a glint of humanity in their eyes. office furniturehomes for sale in tucson

 
At 3/02/2012 11:43:00 PM, Blogger carotkhonglo said...

Richmond is a much sought after residential location, and among the wealthiest areas in the United Kingdom.[3] It is also a significant commercial and upmarket retail centre, and has a developed day and evening economy.inmobiliariassplit pdf

 
At 3/03/2012 05:10:00 AM, Blogger camap said...

The name chosen by the founder of the US city of Richmond, capital of the Commonwealth of Virginia, derives from here. The founder had spent time in Richmond during his youth and knew that the views from the hills overlooking the rivers in both places were similar. Naturally these two Richmonds are twinned.[11] In addition, Richmond is twinned with Fontainebleau, France and Konstanz, Germanyanger management completion letterbathroom partitions

 
At 3/03/2012 05:35:00 AM, Blogger camap said...

Richmond forms part of the Richmond Park UK Parliament constituency. The current Member of Parliament is Zac Goldsmith. Richmond is also part of the South West London Assembly constituency. For elections to the European Parliament it is part of the London constituency.Acer desktop computerLimo Services In Baltimore

 
At 3/03/2012 06:19:00 AM, Blogger camap said...

This park is both a National Nature Reserve and a Site of Special Scientific Interest. It is about three times the size of Central Park in New York and it contains on a permanent basis around 650 red and fallow deer.[14] The park has a number of traffic and pedestrian gates leading to the surrounding areas of Sheen, Roehampton, Putney, Kingston and Ham.entrance matsC Arm tables

 
At 3/03/2012 10:11:00 AM, Blogger camap said...

Two of these are purveyors of fine chocolates: one is The House of Chocolate located on the corner of Brewers Lane, a long-standing family-run business; the other, a stone's throw to the west on Paved Court, is William Curley, a chocolatier whose award-winning fare, together with pastries and hot drinks can be consumed on the premises.Jeep AC CompressorOceanfront homes

 
At 3/05/2012 04:19:00 PM, Blogger trungvkvk said...

Grauer Starcorporate finance

I lay my love on you
It's all I wanna do
Everytime I breathe I feel brand new
You open up my heart
Show me all your love, and walk right through
As I lay my love on you

 
At 3/05/2012 06:50:00 PM, Blogger amantr said...

IT Support Londoncannabis vaporizers
So so I say a little prayer
Hope my dreams will take me there
Where the skies are blue
To see you once again my love
All the seas go coast to coast
Find the place I love the most
Where the fields are green
To see you once again my love
See you in a prayer

 
At 3/07/2012 07:44:00 AM, Blogger carotkhonglo said...

bounded by the Indian Ocean to the west and north. The International Hydrographic Organization (IHO) designates the body of water south of the continent as part of the Indian Ocean, although it is officially gazetted as the Southern Ocean in AustraliaCatering Companies in Atlantalost the weight

 
At 3/07/2012 11:35:00 AM, Blogger mup said...

An exception to this is the northern tropical regions. The Kimberley has an extremely hot monsoonal climate with average annual rainfall ranging from 500 to 1,500 millimetres (20–60 in)Mortgage ProtectionCondos For Sale in Mississauga

 
At 3/07/2012 08:00:00 PM, Blogger carotkhonglo said...

The central four-fifths of the state is semiarid or desert and is lightly inhabited with the only significant activity being mining. Annual rainfall averages 200–250 millimetres (8–10 in), most of which occurs in sporadic torrential falls related to cyclone events in summer months.Jewish educationflats and sandals for 2012

 
At 3/08/2012 05:00:00 AM, Blogger tung said...

Martha Layne Collins (born 1936) is a businesswoman and politician from Kentucky who was the state's 56th governor from 1983 to 1987. Prior to her election as governor, she was the 48th Lieutenant Governor of Kentucky, serving under John Y. Brown, Jr. She is the only woman to have been governor of Kentucky, and her election made her the highest-ranking Democratic woman in the U.S. at the time. She was considered as a possible running mate for Democratic presidential nominee Walter Mondale in the 1984 presidential election, but Mondale chose Geraldine Ferraro instead.
vaporizerscisco routers

 
At 3/09/2012 06:24:00 AM, Blogger carotkhonglo said...

As of 2009 all plans for the mining of the lignite has been suspended indefinitely due to the stock market collapse in early 2009. The sudden drop in oil prices would make the refinement of the oil suspended in the lignite financially unviable.[7]brazilian blowoutpanuveitis

 
At 3/09/2012 07:12:00 AM, Blogger carotkhonglo said...

. At the time Salmon Gums was regarded as being on the fringe of arable land. The low soil fertility and marginal average rainfall of 341mm meant that yields were poor prior to the 1950s. As a result, the land was not in high demand, so the opportunity cost to the government was quite small.Boston Physical TherapyWholesale Womens Shoes

 
At 3/10/2012 01:50:00 AM, Blogger carotkhonglo said...

The southwest coastal area has a Mediterranean climate and was originally heavily forested, including large stands of the karri, one of the tallest trees in the world.[10] This agricultural region of Western AustraliaLocation d\'appartements de vacances à Madèrebed sydney

 
At 3/10/2012 05:45:00 PM, Blogger trungvkvk said...

pokerkingseo company
Like once in a lifetime
You change my world

I lay my love on you
You make me feel brand new
Show me all your love, and walk right through
As I lay my love on you

 
At 3/11/2012 09:45:00 AM, Blogger M.U.P said...

An exception to this is the northern tropical regions. The Kimberley has an extremely hot monsoonal climate with average annual rainfall ranging from 500 to 1,500 millimetres ( bank endorsement stampsinterim management

 
At 3/12/2012 11:05:00 PM, Blogger camap said...

Western Australia's economy is largely driven by extraction and processing of a diverse range of mineral and petroleum commodities. The structure of the economy is closely linked to the abundance of natural resources found in the State, providing a comparative advantage in resource extraction and processingMiddle East Classified Ads send text

 
At 3/13/2012 12:57:00 PM, Blogger 12345 said...

Wind Energy Tax Credits
Lancaster Web DesignTitan's ionosphere is also more complex than Earth's, with the main ionosphere at an altitude of 1,200 km but with an additional layer of charged particles at 63 km. This splits Titan's atmosphere to some extent into two separate radio-resonating chambers. The source of natural ELF waves (see Bulk characteristics above) on Titan is unclear as there does not appear to be extensive lightning activity.

 
At 3/14/2012 01:46:00 AM, Blogger camap said...

Europeans began to settle permanently in 1826 when Albany was claimed by Britain to forestall French claims to the western third of the continent. Perth was founded as the Swan River Colony in 1829 by British and Irish settlersenterprise document management software Small Business Web Designer

 
At 3/14/2012 10:09:00 AM, Blogger camap said...

though the outpost languished, eventually requesting convict labour to augment its population. In the 1890s, interstate migration resulting from a mining boom in the Goldfields region resulted in a sharp population increase. schoolbordfolie CAD

 
At 3/17/2012 03:17:00 AM, Blogger camap said...

The Great Egret is a large bird with all-white plumage that can reach one meter in height, weigh up to 950 grams (2.1 lb) and a wingspan of 165 to 215 cm (65 to 85 in). It is thus only slightly smaller than the Great Blue or Grey Heron (A. cinerea).Igre gold charms for bracelets

 
At 3/18/2012 03:08:00 AM, Blogger camap said...

. He was the great-grandson of William Stevenson, brother of Vice President Adlai E. Stevenson, making him a second cousin once removed of presidential candidate Adlai E. Stevenson II.affordable Powered wheelchair Wholesale Gowns

 
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...

Tesco Direct Discount CodesProactol Plus Reviews
When does my "someday" begin
When I'll find someone again
And what if I still am not truly over
What am I supposed to do then, babe...
Do you see what I'm saying
Even if, if it's not making sense
So when can I see you

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

psicologos villanueva pardillo

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...

psicologia clinica las rozas

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.
get a slut

Decoding

 
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.
psicologia clinica aravaca

queda de cabelos

 
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.
web design leeds

jobs schweiz

 
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
stainless steel fabrication

top view men

 
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.
Chiropractors in Fort Collins

Audiolibros Gratis

 
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.
Get Rid of Stretch Marks

medicare supplements texas

 
At 4/28/2012 09:49:00 AM, Blogger HT said...

Cotton Candy Maker Machine

Descargar Programas Gratis
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...

Restaurant Text Message Marketing

itunes card instant
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...

caravan annexesAuto Warranty
I'm trembling inside
And nobody knows it but me (yeah)

Lie awake, it's a quarter past three
I'm screaming at night if I thought you'd hear me
Yeah, my heart is calling you
And nobody knows it but me (well, well)

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

balers

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...

Credit Card Databasedomain anmelden
If you search for tenderness
it isn't hard to find.
You can have the love you need to live.
But if you look for truthfulness
You might just as well be blind.
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.
small pet carrier

enagic distributor

 
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.
water ionizer

yiwu china

 
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.
Travel Photographer

Baja Homes For Sale

 
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.
Restaurant Hardware

Rosarito Condos

 
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.
braces orlando

xanthan gum food grade

 
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.
benefits of alkaline water

you are a winner

 
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.
humourous illustration

jobs

 
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.
Burnaby Real Estate

Land Rover HID

 
At 5/04/2012 02:55:00 PM, Blogger trung said...

engine treatmenttruck driver
For what is a man, what has he got?
If not himself, then he has naught.
To say the things he truly feels;
And not the words of one who kneels.
The record shows I took the blows -
And did it my way!uct

 
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.
water ionizer

تخسيس الوزن

 
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.
water ionizer

Tomball Chiropractic

 
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
torno cnc

estetica y belleza las rozas

 
At 5/05/2012 04:24:00 PM, Blogger trung said...

Bald Eagle picturestop ten lists
I miss you standing by my side
I'm dying here its clear to see
There ain't no you, God knows there ain't no me
Don't want to live, I want to die
If I can't have you in my life

 
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!
börse dax

Saskatoon apartments for rent

 
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.
photo decoration interieur

memory sticks

 
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.
Spyderco

coupe de cheveux homme

 
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.
cabo fitness

electric fireplace heaters

 
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.
brain evolution system

g-20 los cabos

 
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
cabo san lucas

make money from home

 
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.
intercom installation

forex trade rebates

 
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.
Milton Homes For Sale

microsoft office 2010 product key

 
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.
hot tubs

Paid Work Abroad

 
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!
powerpoint search

vietnam voyages

 
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,
nhac cho vietnamobile

Doładowanie telefonu

 
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.
made to measure suits

Promotional Coasters

 
At 11/25/2013 06:05:00 PM, Blogger Jay Lamptey said...

I remember talking to the agent John Jarrold and he mentioned how, in his editorial days, he used to like taking the one positive line from a bad review
http://www.pure-jobs.com

 

Post a Comment

Links to this post:

Create a Link

<< Home