text:
(born 1951). American actress Anjelica Huston was noted for her portrayal of strong self-sufficient women. She won an Academy Award for best supporting actress for her role as a Mafia daughter in the comedy drama Prizzi's Honor (1985).
Huston was born on July 8, 1951, in Los Angeles, California. Her father was film director John Huston, and her grandfather was actor Walter Huston. In 1953 her family moved from California to Ireland. John Huston was absent for long periods while he worked on film sets. In 1962 Anjelica moved with her mother and brother to London, England, and eventually became involved in political activism.
Huston left school in 1968 and subsequently auditioned for a role in Franco Zeffirelli's movie Romeo and Juliet. Instead, however, her father cast her in his production of A Walk with Love and Death (1969). The period romance was panned, as was Huston's performance. She turned to theater and furthered a modeling career. A 1980 car crash reignited her desire to act. She subsequently took acting lessons and appeared in supporting roles, including in the film The Postman Always Rings Twice (1981) and on the television show Laverne & Shirley in 1982–83.
For her role in the 1985 film Prizzi's Honor, which was directed by her father, Huston earned an Oscar for best supporting actress. In 1987 she played a woman cajoled into revealing dark secrets in The Dead, which was also directed by her father. Other films from the 1980s in which Huston appeared include the dramas Gardens of Stone (1987), A Handful of Dust (1988), and Crimes and Misdemeanors (1989).
Huston earned two more Academy Award nominations for her portrayal of a Holocaust survivor who finds that her husband has remarried in Enemies: A Love Story (1989) and for her role as a murderous con artist in The Grifters (1990). She then played a series of matriarchs in movies such as The Addams Family (1991) and its sequel, Addams Family Values (1993), The Perez Family (1995), Buffalo '66 (1998), and Ever After: A Cinderella Story (1998). In 1996 Huston directed Bastard out of Carolina, a cable movie, and in 1999 she directed and starred in the film Agnes Browne.
Huston continued to act in the 21st century in films such as The Royal Tenenbaums (2001), The Life Aquatic with Steve Zissou (2004), The Darjeeling Limited (2007), Choke (2008), and 50/50 (2011). In 2012–13 she appeared on the television series Smash. Huston related episodes from her early life in A Story Lately Told: Coming of Age in Ireland, London, and New York (2013).


Thank you to everyone who supported the third annual Patter Merchants Comedy Clash...What a night!
Our host Capital FM's Des Clarke was on top form and kept all of our acts in check - huge thanks to the hilarious Jamie Dalgleish, Megan Shandley, Christopher Macarthur-Boyd and the legend "Big" Gary Little.
With thanks to all of our sponsors and prize donors and of course our table hosts and guests on the night for your generosity - we raised over £12,500!
Keep an eye out for the 2020 date to be announced soon!


text:
Oil & Gas / North Sea
Wood wins £58million Mariner contract with Equinor, supporting 100 jobs
Wood has secured a $75million (£58.2m) contract with Equinor for its Mariner field, supporting around 100 jobs.
14/10/2020, 8:04 am Updated: 14/10/2020, 1:40 pm
© Abermedia / Michal Wachucik
The Aberdeen-headquartered energy services firm will provide operations, maintenance, modifications and offshore services to the Mariner A platform and Mariner B floating storage unit over three years.
Wood said the deal will run from January 2021 to Q4 2023, with options to extend.
It comes after Equinor started up the multi-billion pound development east of Shetland last year, which is expected to continue producing through to 2050.
Craig Shanaghey, president of Wood's operations services business in Europe and Africa, said: "We are delighted to extend our strong partnership with Equinor to include support for their operations at the pioneering Mariner field.
"Wood has a long-standing track record of partnering with our clients to deliver safe, reliable, and optimised operations in the UKCS, and we look forward to extending that to include Mariner by leveraging our deep operational knowledge, experience, and digital capability.
"Mariner is still in its early years of production and, with Wood's ambition to realise a digitally-enabled future, we see excellent potential to explore new opportunities that will promote a lifetime of sustainable and responsible operations at the field."
The work will support a mix of around 100 onshore and offshore roles, Wood said, delivered by teams based out of Aberdeen.
It comes as the firm announced in August it had reduced headcount by 10,000 jobs roles globally since the downturn, including some in north-east Scotland.
Mariner is Equinor's first operated asset in the UK, with its next target being Rosebank to the west of Shetland, though a final investment decision on that project is not expected until 2022.
The development has been praised for its use of automated drilling and digital twin technologies to improve costs.
This latest contract builds on Wood's agreements with Equinor on the Kollsnes gas processing facility and Breidablikk tie-back development in Norway.
Borr Drilling appoints CFO who was replaced amid Weatherford bankruptcy
Craig Shanaghey
East of Shetland
NEO Energy enters exclusive talks with ExxonMobil over North Sea portfolio
EnQuest wraps up deal for stake in Bressay field
Energean hits go on NEA/NI project

redpajama_set_name: RedPajamaCommonCrawl


text:
extra_balls:
============

*Config file section*

+----------------------------------------------------------------------------+---------+
| Valid in :doc:`machine config files </config/instructions/machine_config>` | **NO**  |
+----------------------------------------------------------------------------+---------+
| Valid in :doc:`mode config files </config/instructions/mode_config>`       | **YES** |
+----------------------------------------------------------------------------+---------+

.. overview

The ``extra_balls:`` section of your config is where you configure
which events trigger and reset extra ball awards.

Note that this extra ball abstract device only takes care of awarding the
extra ball and can lit a light when enabled/available.
The logic to qualify for an extra ball has to be implemented in your mode.
See :doc:`/game_logic/extra_balls/index` for more details.

Here's an example:

.. code-block:: mpf-config

   ##! mode: mode1
   extra_balls:
     my_mode_eb:
       award_events: alien_smashed

In the above example, the extra ball called ``my_mode_eb`` will be
given to the player when the event ``alien_smashed`` is posted. After that,
future ``alien_smashed`` events will not lead to additional extra balls. (The
``my_mode_eb`` extra ball is "used up", in a sense).

This is all tracked per-player in a player variable dictionary called "extra_balls_awarded"

.. config

Optional settings
-----------------

The following sections are optional in the ``extra_balls:`` section of your config. (If you don't include them, the default will be used).

award_events:
~~~~~~~~~~~~~
List of one (or more) device control events (:doc:`Instructions for entering device control events </config/instructions/device_control_events>`). Defaults to empty.

Events in this list, when posted, award this extra ball to the current player.

enabled:
~~~~~~~~
Single value, type: ``boolean`` (``true``/``false``). Default: ``true``

Whether the device starts enabled or disabled.

group:
~~~~~~
Single value, type: string name of a :doc:`extra_ball_groups <extra_ball_groups>` device. Defaults to empty.

The extra ball group which this ball belongs to which can further limit the
maximum number of balls and enable/disable the device.

light_events:
~~~~~~~~~~~~~
List of one (or more) device control events (:doc:`Instructions for entering device control events </config/instructions/device_control_events>`). Defaults to empty.

Event to light the extra ball (if enabled).

max_per_game:
~~~~~~~~~~~~~
Single value, type: ``integer``. Default: ``1``

Maximum number of extra balls to award per player for this particular extra
ball device.
This might be further limited by the extra_ball_group max_per_game limit.
In that case if either of the two limits is exceeded no more balls will be
awarded.

console_log:
~~~~~~~~~~~~
Single value, type: one of the following options: none, basic, full. Default: ``basic``

Log level for the console log for this device.

debug:
~~~~~~
Single value, type: ``boolean`` (``true``/``false``). Default: ``false``

See the :doc:`documentation on the debug setting </config/instructions/debug>`
for details.

file_log:
~~~~~~~~~
Single value, type: one of the following options: none, basic, full. Default: ``basic``

Log level for the file log for this device.

label:
~~~~~~
Single value, type: ``string``. Default: ``%``

A descriptive name for this device which will show up in the service menu
and reports.

tags:
~~~~~
List of one (or more) values, each is a type: ``string``. Defaults to empty.

Special / reserved tags for extra balls: *None*

See the :doc:`documentation on tags </config/instructions/tags>` for details.

Related How To guides
---------------------

* :doc:`/game_logic/extra_balls/index`

redpajama_set_name: RedPajamaGithub


text:
'Bank of Mum and Dad' set to finance 25% of all UK mortgages in 2016
Home / News / 'Bank of Mum and Dad' set to finance 25% of all UK mortgages in 2016
The so-called 'Bank of Mum and Dad' are now as crucial as one of the UK's top 10 lenders – and will finance 25 per cent of all mortgages in in 2016.
That is according to research from Legal & General and financial services group Cebr.
Their research has indicated that parents will lend over £5 billion this year, providing deposits for over 300,000 mortgages and purchasing homes worth £77 billion.
The Bank of Mum and Dad's average financial contribution is £17,500, or seven per cent of the average purchase price of a home.
Nigel Wilson, the CEO of Legal & General has said that parental support is becoming increasingly important to help young people get on the housing ladder.
And he warned : "Relying so heavily on the bank of mum and dad however risks increasing inequality, as many young people today are not lucky enough to be able to access parental support when buying a home, or can't afford to buy even with parental help.
"We have a supply-side problem in housing – we are simply not building enough houses. We need to build more, especially as the Bank of Mum and Dad could soon start to experience a funding crisis of its own."
The research also showed that three-quarters of family-funded house purchases are done by parents, with family friends and grandparents also making up the difference.
Episode 20 of Lifetime's Desert Island Discs – featuring Office Administrator Mathew Leech
Teams gearing up for 2016 Lifetime-sponsored Barnsley Hospice Golf Day


Make your contact forms user friendly and fast. Your customers will not wish to wait until the page reloads after form submission. And with "Extended Contact Form" module for Prestashop they won't have to. Also your customers will see all the necessary information about your store on contact page.


text:
Institution: Massachusetts Inst of Technol MIT Libs
Early Career Reviewers
FlyBook
WormBook
YeastBook
GSA Honors and Awards
Methods, Technology & Resources
Toolbox Reviews
Effects of trans-acting Genetic Modifiers on Meiotic Recombination Across the a1–sh2 Interval of Maize
Marna D. Yandeau-Nelson, Basil J. Nikolau and Patrick S. Schnable
Genetics September 1, 2006 vol. 174 no. 1 101-112; https://doi.org/10.1534/genetics.105.049270
Marna D. Yandeau-Nelson
Basil J. Nikolau
Patrick S. Schnable
Meiotic recombination rates are potentially affected by cis- and trans-acting factors, i.e., genotype-specific modifiers that do or do not reside in the recombining interval, respectively. Effects of trans modifiers on recombination across the ∼140-kb maize a1–sh2 interval of chromosome 3L were studied in the absence of polymorphic cis factors in three genetically diverse backgrounds into which a sequence-identical a1–sh2 interval had been introgressed. Genetic distances across a1–sh2 varied twofold among genetic backgrounds. Although the existence of regions exhibiting high and low rates of recombination (hot and cold spots, respectively) was conserved across backgrounds, the absolute rates of recombination in these sequence-identical regions differed significantly among backgrounds. In addition, an intergenic hot spot had a higher rate of recombination as compared to the genome average rate of recombination in one background and not in another. Recombination rates across two genetic intervals on chromosome 1 did not exhibit the same relationships among backgrounds as was observed in a1–sh2. This suggests that at least some detected trans-acting factors do not equally affect recombination across the genome. This study establishes that trans modifier(s) polymorphic among genetic backgrounds can increase and decrease recombination in both genic and intergenic regions over relatively small genetic and physical intervals.
MEIOTIC recombination mediates the proper disjunction of chromosomes, generates novel allelic combinations across chromosomes, and provides the genetic diversity upon which selection can act. According to the double-strand break repair (DSBR) model (Szostak et al. 1983; Sun et al. 1991), meiotic recombination initiates with a double-strand break (DSB). Fungal, animal, and plant chromosomes exhibit regions of recombination hyperactivity, i.e., hot spots, and hypoactivity, i.e., cold spots (reviewed in Lichten and Goldman 1995; Schnable et al. 1998; Petes 2001; Nachman 2002), as compared to the genomewide average rate of recombination in the organism being studied. Consistent with the DSBR model, recombination hot spots in Saccharomyces cerevisiae are clearly associated with DSBs (reviewed in Lichten and Goldman 1995) and these DSB hot spots are not distributed randomly across the yeast genome (Gerton et al. 2000).
Nonrandom patterns of recombination have been identified in all organisms studied to date. For example, in most species recombination rates per megabase are suppressed in and surrounding centromeres (Lambie and Roeder 1986; Werner et al. 1992; Mahtani and Willard 1998; Puechberty et al. 1999; Kagawa et al. 2002). Similarly, in most species, with the exception of Caenorhabditis elegans, for which recombination rates are negatively correlated with gene density (Barnes et al. 1995), recombination rates are higher in gene-rich than in gene-poor regions of the genome (reviewed in Lichten and Goldman 1995; Schnable et al. 1998). In wheat and barley, for example, analyses of integrated genetic and cytogenetically based physical maps (Gill et al. 1993, 1996a,b; Hohmann et al. 1994; Delaney et al. 1995a,b; Mickelson-Young et al. 1995; Kunzel et al. 2000) have shown that most recombination occurs in relatively small gene-rich regions. Similarly, in maize recombination nodules cluster in regions of high gene density (Anderson et al. 2006). In addition, it is estimated that recombination in a gene-dense portion of the bz1 region on chromosome 9S in maize is two orders of magnitude greater than in a flanking region of high retrotransposon density (Fu et al. 2002).
Although it is well established that high frequencies of recombination are positively correlated with gene-rich regions in the grasses, recombination breakpoints within these hot spots might resolve equally in genic and intergenic sequences. A high-resolution recombination mapping study across the ∼140-kb multigenic maize a1–sh2 interval supports that genes per se are preferred recombination hot spots and intergenic regions are cold spots; even so, nongenic hot spots and genic cold spots do exist (Yao et al. 2002). These maize genic hot spots can have either nonrandom distributions of breakpoints as observed in a1 (Xu et al. 1995) and other genes (Eggleston et al. 1995; Patterson et al. 1995) or uniform distributions across the gene, as in bz1 (Dooner and Martinez-Ferez 1997).
Variations in meiotic recombination rates occur not only among various regions of the genome but also among genetic backgrounds (reviewed in Simchen and Stamberg 1969) as evidenced in both animals (Koehler et al. 2002) and plants. In Arabidopsis, recombination rates vary among accessions (Sanchez-Moran et al. 2002). In maize, heterogeneity in recombination frequencies has been well documented within corn belt (Beavis and Grant 1991; Tulsieram et al. 1992), corn belt × exotic germplasm (Tulsieram et al. 1992), synthetic (Fatmi et al. 1993), exotic, and teosinte-maize hybrid (Williams et al. 1995) mapping populations and has often confounded the generation of composite genetic maps. Recombination rates in intervals with heterogenous frequencies varied two- to threefold among diverse mapping populations (Williams et al. 1995). Within a mapping population, recombination frequencies both in adjacent (Tulsieram et al. 1992) and between genetically unlinked (Beavis and Grant 1991; Fatmi et al. 1993) regions were often not correlated, demonstrating that recombination can be differentially regulated across a chromosome. But in addition, several intervals on different chromosomes were shown to have either negatively or positively correlated recombination frequencies (Fatmi et al. 1993). Together, these studies suggest that the control of variation in recombination rates might be polygenic and that some factors might control specific genetic intervals.
The heterogeneity in recombination frequencies among genetic backgrounds can be attributed to two general classes of genetic factors: cis and trans. Cis-acting elements are genetic factors that affect recombination in the region in which the factor resides. A study of recombination across several teosinte a1–sh2 intervals, each having large insertion/deletion polymorphisms (IDPs) as compared to maize, has revealed significant differences in recombination rates and distribution of recombination breakpoints across the interval caused by the action of cis factors (Yao and Schnable 2005). Similarly, a not yet molecularly characterized cis factor increases recombination in the large A188 Sh1–Bz1 interval on maize chromosome 9S (Timmermans et al. 1997).
In contrast to cis elements, trans-acting factors are genetic modifiers that are not closely linked to the interval in which they affect recombination. Trans modifiers might include factors involved in chromatin remodeling, proteins involved in the recombination machinery, and autonomous transposons. For example, the rate of crossover (CO) increases near a Mu insertion in a1 in the presence of MuDR transposase (Yandeau-Nelson et al. 2005). Also, recombination is affected on the maize chromosome 9 c1–sh1 and bz1–wx1 intervals as well as chromosome 7 by a single unidentified trans factor (Timmermans et al. 1997). The maize mapping studies described earlier established recombination rate heterogeneity among genetic backgrounds and suggest that this variation is under polygenic control (i.e., multiple trans factors). However, these studies (Tulsieram et al. 1992; Fatmi et al. 1993) were unable to distinguish between the effects of cis- and trans-acting genetic modifiers, both of which potentially influence recombination rates. Also, available data only suggest that rates of recombination differ among genetic backgrounds, but the manner in which factors can affect the distribution of recombination breakpoints across a small physical distance containing both genic and nongenic intervals is unknown.
To elucidate the effects of trans-acting modifiers specifically, these factors must be studied in the absence of polymorphic cis-acting effects. This can be achieved by studying recombination across an interval that is sequence identical in several different genetic backgrounds. The ∼140-kb a1–sh2 interval on maize chromosome 3L is an ideal system in which to study how trans factors modulate both recombination rates and the distribution of recombination breakpoints across an interval because (1) recombination events across the interval are easily identified by nonparental kernel phenotypes, (2) high-resolution mapping of recombination breakpoints is straightforward due to a significant degree of sequence polymorphism between the A1 Sh2 haplotypes and the availability of previously developed IDP markers across much of the interval (Xu et al. 1995; Yao et al. 2002), and (3) the interval is multigenic (Yao et al. 2002), which allows for the study of how trans-acting factors differentially affect recombination across genic and nongenic regions.
A sequence-identical a1∷rdt sh2 interval that had been introgressed into three unique genetic backgrounds (maize inbreds A632, Oh43, and W64A) was used to assess the extent to which trans-acting genetic modifiers that were polymorphic among the backgrounds affect meiotic recombination in the ∼0.1-cM a1–sh2 interval. This study reveals that trans-acting factors can influence both rates of recombination and distribution of recombination breakpoints within conserved hot and cold spots and can convert an average spot to a hot spot. The modifier(s) affecting recombination across this small interval on chromosome 3L does not appear to differentially affect recombination among genetic backgrounds in the same way across an interval on chromosome 1S, suggesting that at least some trans-acting factors affect specific regions of the genome.
Genetic stocks:
The shrunken-2 (sh2) gene is located on chromosome 3 ∼0.1 cM centromere-distal to the a1 locus (Civardi et al. 1994). Mutations at these loci condition shrunken and colorless kernel phenotypes, respectively. The a1∷rdt sh2 haplotype (GenBank accession no. AF072704) was described previously (reviewed in Xu et al. 1995). The A1-LC Sh2 haplotype (GenBank accession nos. AF434192, AF347696, AF363390, X05068, and AF363391) is derived from the inbred line C (LC), a color-converted version of W22.
The a1∷rdt sh2 (A632)7, a1∷rdt sh2 (Oh43)9, and a1∷rdt sh2 (W64A)6 genetic stocks were gifts from David Glover (Purdue University). To generate these stocks, the inbreds A632, Oh43, and W64A were crossed by a line that was homozygous for the a1∷rdt sh2 haplotype. Using the a1 and sh2 mutants as markers, the a1∷rdt sh2 haplotype was backcrossed into the inbreds for seven, nine, and six generations to generate the trans stocks a1∷rdt sh2 (A632)7, a1∷rdt sh2 (Oh43)9, and a1∷rdt sh2 (W64A)6, respectively. The trans stocks were self-pollinated to generate homozygous a1∷rdt sh2 sources in each of the three genetic backgrounds.
Isolation of genetic recombinants and estimations of genetic distances:
Recombinants across a1–sh2:
To identify recombinants that resolve between a1 and sh2, each of the three inbred trans stocks was used as a male onto the inbred line C (cross 1). Recombination was measured in the three resulting trans stock/line C F1's, i.e., A632/LC, Oh43/LC, and W64A/LC, via testcrosses (cross 2) as described by Yao et al. (2002) and Yao and Schnable (2005). To control environmental effects on recombination, these testcrosses were performed during a single season (summer, 1997) in a single field.
Cross 1: A1-LC Sh2/A1-LC Sh2 (line C) × a1∷rdt sh2/a1∷rdt sh2 (trans stock)
Cross 2: A1-LC Sh2/a1∷rdt sh2 (F1 from cross 1) × a1∷rdt sh2/a1∷rdt sh2
Progeny from cross 2 segregated for parental colored round and colorless shrunken kernels. Rare kernels with nonparental phenotypes (viz., colored shrunken and colorless round) putatively carry recombination events between a1 and sh2. Because the original A632, Oh43, and W64A inbreds are recessive for r1, a gene that encodes a transcription factor that acts upstream of a1 in anthocyanin biosynthesis, colorless round (a1′Sh2) kernels could be generated due to the absence of r1 and not because of resolution of a recombination breakpoint between a1 and sh2. For this reason, only the colored shrunken (A1′sh2) recombinant class was analyzed in this study. Putative A1′sh2 recombinant kernels from each inbred background were tested, confirmed, and made homozygous as described in Xu et al. (1995). The actual number of A1′sh2 recombinants isolated from each genetic background was estimated from the frequency of confirmed recombinants out of the total number analyzed.
Recombinants across chromosome 1S:
To identify genetic intervals unlinked to the a1–sh2 interval on chromosome 3L, ∼1000 IDP genetic markers developed by the Maize Genetic Mapping Project (http://maize-mapping.plantgenomics.iastate.edu/) and genetically mapped to chromosomes 1–2, and 4–10 were surveyed. The criteria for selecting genetic intervals were that: (1) they be defined by two IDP markers that were 1–20 cM apart, (2) each IDP marker was polymorphic between line C (the A1 Sh2 haplotype used in cross 1) and the three trans stocks, and (3) the genetic interval could be assayed in each of the three trans-stock experiments. Only two intervals that met these criteria were identified; both were located on chromosome 1S. Interval 1S.1 is defined by genetic markers IDP194 and IDP254 and 1S.2 is defined by markers IDP112 and IDP643. The IDP194 and IDP254 markers were designed from MEST54-C06 (GenBank accession no. BM072826) and MEST139-B10 (GenBank accession no. BM334583), respectively, and map to positions 115.6 and 115.0 cM, respectively, on the maize IDP_body map version 4 (http://magi.plantgenomics.iastate.edu/cgi-bin/cmap). Markers IDP112 and IDP643 were designed from MEST19-B03 (GenBank accession no. BG841229) and MEST129-G08 (GenBank accession no. BM333984) and map to positions 82.8 and 84.6 cM, respectively.
Mapping populations were generated by crossing a single F1 plant from cross 1 for each of the trans stocks to the inbred B77 (cross 3).
Cross 3: B77 × A1-LC Sh2/a1∷rdt sh2 (F1 from cross 1)
Consequently, the progeny of cross 3 potentially carry alleles from the specific trans stock, line C, and B77. B77 was selected as the female parent for cross 3 because when tested with a sample of IDP markers across the genome it showed a high frequency of polymorphisms relative to line C and each of the three trans stocks (data not shown). Recombination rates across 1S.1 and 1S.2 were assayed in populations of ∼700 seedlings derived from progeny of cross 3 in each genetic background. Recombinants across each interval were identified as those individuals having nonparental combinations of genotypes at the two linked loci in each interval. Genetic distances in each background were calculated using the total number of recombinants identified in each trans stock and the total number of seedlings analyzed.
Genotypes were determined using PCR primers IDP112F, 5′-CTGTGACATGTTTGATGCCC-3′, IDP112R, 5′-GGTGATGACCACGTACAAGC-3′, and IDP112AW, 5′-CCC TGC TGA TAG TGA TAG AC-3′; IDP643F, 5′-ACCCTCATCTTCAGCAG TCG-3′ and IDP643R, 5′-GGTGAAACGGCAGTACAAGG-3′; IDP194F, 5′-GACAGATCCCTAACACTTGGG-3′ and IDP194R, 5′-AACAAGGCAACCTGTGAAGC-3′; and IDP254F, 5′-ATGTTGGTTGAGCCTCTTGG-3′ and IDP254R, 5′-GATTCAGAGAGAGTGCATGGC-3′.
Seed sterilization, germination, and DNA isolation:
Colored shrunken kernels that were homozygous for recombinant A1′sh2 haplotypes originally isolated from cross 2 and ∼700 kernels from each genetic background derived from cross 3 were germinated in 96-well flats. PCR-ready DNA was isolated as described (Dietrich et al. 2002).
Due to fungal contamination, many shrunken kernels do not germinate in the soil. For these recombinants, a second aliquot of shrunken kernels was sterilized with pure bleach (Clorox) for 1 min and then rinsed repeatedly (10–15 times) with sterile water. Sterilized kernels were plated on moistened autoclaved germination paper (Anchor Paper, St. Paul) in sterile Petri dishes, treated with 0.5% Captan 50-W fungicide (Platte Chemical, Fremont, NE), and covered with activated charcoal. Seeds were germinated at 28° for 5–7 days. Seeds were transferred to new plates, watered, and covered with fresh charcoal every 1–2 days. Tissue was collected from coleoptiles and DNA isolation was as described above.
Physically mapping recombination breakpoints:
Recombination breakpoints across the a1–sh2 interval were mapped relative to eight IDP markers that had previously been identified between the A1-LC Sh2 and a1∷rdt sh2 haplotypes (YAO et al. 2002). These markers separated the a1–sh2 interval into eight informative subintervals (Figure 1A); intervals are numbered according to Yao et al. (2002). Allele-specific primers not described by Yao et al. (2002) are listed below. Primers that when paired recognize a size polymorphism between the two haplotypes are XL3, 5′-ATGAGCGGGAGCCTATG-3′, and XL4, 5′-TCAGCATCCATACCATTG-3′. Allele-specific and size polymorphism primers are shown in Figure 1A.
Distributions of recombination breakpoints across the a1–sh2 interval in the A632/LC, Oh43/LC, and W64A/LC genetic backgrounds. (A) A schematic of the A1-LC Sh2 and a1∷rdt sh2 haplotypes in which boxes represent genes. Subintervals are defined by allele-specific PCR primers designed at sequence polymorphisms that distinguish the haplotypes and are numbered according to Yao et al. (2002). Allele-specific primers are indicated by arrows above the haplotype they specifically amplify. The size of each subinterval in the line C haplotype is listed. As compared to Yao et al. (2002), subintervals XI, XII, and XIII have been combined into a single interval (XI–XIII). Because subintervals IV and V are very small and no recombinants mapped to these subintervals, they are considered not informative in this study. (B) Rates of recombination in each subinterval. The * and ** denote significant differences relative to the genomewide average recombination rate at the 0.05 and 0.01 levels, respectively. The rates of recombination in subintervals IX and XI–XIII are so low that they are not visible on this graph. Subinterval X contains the 3′ portion of x1. Even if all of the recombinants that resolved within subintervals XI–XIII resolved within x1, the rate of recombination in x1 would still be significantly lower than the genomewide average in each of the three backgrounds (P-value < 0.002). (C) The percentage of recombination breakpoints that mapped to each subinterval. (D) The number of recombination breakpoints that mapped to each subinterval. (C and D) In some cases, no breakpoints mapped to a subinterval in a given genetic background and therefore no bar is shown. (B–D) Data from Yao et al. (2002) are provided for reference.
The breakpoints associated with >85% of the confirmed recombinants from the A632 (90%; 117/130), Oh43 (87%; 155/178), and W64A (85%; 185/219) genetic backgrounds were physically mapped. It was not possible to map the remainder due to difficulties in obtaining homozygous recombinant chromosomes or in germinating shrunken kernels.
Statistical methods:
For each physical interval examined, genetic distances (in centimorgans) were compared among genetic backgrounds using χ2 homogeneity tests. For the genetic distances between a1 and sh2, the corrected number of A1′sh2 recombinants (Table 1) was doubled because the reciprocal class, a1′Sh2, was not analyzed. This analysis makes the reasonable assumption that the rates at which the two recombinant classes arise are equal.
Genetic distances across a common a1–sh2 interval in three genetic backgrounds
Statistical methods and population size adjustments were performed as described by Yao and Schnable (2005). The actual numbers of mapped recombinants and the adjusted population sizes were used for subsequent χ2 homogeneity tests. Pearson χ2 tests were conducted as described by Yao and Schnable (2005) to compare the actual recombinant breakpoint distribution in each genetic background across a1–sh2 to (1) the expected distribution if recombination across the interval was equal to the genomewide average (2.4 cM/Mb) and (2) the expected distribution of recombinants if the rate of recombination across each subinterval was equal to the average recombination rate across the entire a1–sh2 interval. In this study, rates of recombination were compared to a genomewide average recombination rate of 2.4 cM/Mb (Table 2). This is an updated rate compared to the 2.1 cM/Mb rate (YAO et al. 2002) based on a more recent maize genetic map that consists of 5917 cM (Lee et al. 2002).
Statistical analyses of recombination across a1–sh2
The expected numbers of recombinants for each subinterval were calculated using the subinterval distances measured in the line C haplotype (Figure 1A). The sizes of each subinterval were previously determined via sequencing or DNA gel blot analyses (Yao et al. 2002). For the a1∷rdt sh2 haplotype, interval sizes were determined using the genomic sequence from a1 to yz1 of this haplotype (GenBank accession no. AF072704) for subintervals II (1.7 kb), III (1.34 kb), IV (0.4 kb), V (80 bp), VI (2.2 kb), VII (620 bp), and VIII (3.7 kb). To test if the use of subinterval sizes from line C affects comparisons of recombination rates to the genomewide and to the a1–sh2 interval average rates of recombination, the expected numbers of recombinants and the corresponding statistical analyses were also conducted using subinterval sizes from the a1∷rdt sh2 haplotype. Because the significance or nonsignificance for each subinterval was not altered by which haplotype was used to determine sizes of each subinterval, only statistical analyses using subinterval sizes measured in line C are presented.
To investigate which subintervals contribute to the different breakpoint distributions among the backgrounds, an exploratory statistical analysis was conducted. Freeman–Halton tests as described in Yao and Schnable (2005) were conducted in which each subinterval was successively removed (and then replaced) from the analysis. If the recombination rates in a subinterval were contributing to differences in breakpoint distributions across the entire interval, then upon removal of this subinterval (or group of subintervals) the backgrounds should no longer differ significantly.
The genetic distance between a1 and sh2 and the distribution of recombination breakpoints varies among genetic backgrounds:
A common a1∷rdt sh2 interval was introgressed into the inbred lines A632, Oh43, and W64A for multiple generations (materials and methods). Each of the resulting trans stocks contains a sequence-identical a1∷rdt sh2 interval on chromosome 3L, while the bulk of its genome is expected to be derived from the recurrent parent: i.e., A632, Oh43, and W64A. These stocks are ideal for studying the effects of trans-acting genetic modifiers on recombination across the a1–sh2 interval because the presence of an identical a1–sh2 interval in each of the three backgrounds ensures that the cis factors that affect recombination within the a1–sh2 interval are identical in each of the inbred backgrounds. To analyze the effects of genetic background on recombination across the a1–sh2 interval, each of the trans stocks was crossed to the inbred line C. Because each of the resulting F1's is identically heterozygous at the a1–sh2 interval (A1-LC Sh2/a1∷rdt sh2), the only modifiers of recombination that are expected to differ among these three F1's should be outside of the a1–sh2 interval. Hence, any observed differences in meiotic recombination across the a1–sh2 interval among the three genetic backgrounds can be attributed specifically to trans factors.
Meiotic recombination events that resolved between a1 and sh2 and yielded colored shrunken kernels (A1′sh2) were identified from cross 2 (materials and methods). Colorless round (a1′Sh2) kernels were not analyzed for reasons described in materials and methods. Each of the genetic backgrounds exhibited a statistically different rate of recombination across the a1–sh2 interval and these rates varied almost twofold (Table 1).
Rates of recombination across the a1–sh2 interval differ significantly due to the action of trans-acting modifier(s). To determine whether these trans-acting factors also affect the distributions of meiotic recombination breakpoints across genic and nongenic regions in this interval, recombination breakpoints isolated from each genetic background were mapped relative to sequence polymorphisms between the A1-LC Sh2 and a1∷rdt sh2 haplotypes (materials and methods). Because the a1–sh2 intervals present in each of the genetic backgrounds were identical, breakpoints could be mapped relative to the same sequence polymorphisms, thereby facilitating comparisons among genetic backgrounds.
In each background recombination breakpoints were not randomly distributed across the a1–sh2 interval (P < 4.0e−39). Indeed, >95% of the recombinants isolated from each background mapped within the a1–yz1 interval (subintervals II–VIII; Figure 1, C and D) that composes only ∼10% of the physical length of a1–sh2. The majority of the remaining recombinants resolved at the 3′ end of x1 (subinterval X; Figure 1, C and D). These observations are consistent with previously described distributions of recombination breakpoints across the a1–sh2 interval (Yao et al. 2002; Yao and Schnable 2005).
The distributions of recombination breakpoints across the eight a1–sh2 subintervals defined by sequence polymorphisms between the A1-LC Sh2 and a1∷rdt sh2 haplotypes (Figure 1) were compared among genetic backgrounds (materials and methods). The distribution of breakpoints in the Oh43/LC background differed significantly from the distributions in both the A632/LC and W64A/LC backgrounds (Table 3). This finding indicates that trans-acting factors that are polymorphic in the Oh43/LC background as compared to the other two genetic backgrounds affect not only rates of recombination across the entire a1–sh2 interval but also the distribution of recombination breakpoints within this interval. Using the exploratory statistical analysis described in materials and methods it was possible to establish that in the Oh43/LC background the interloop (subinterval VI) and the yz1 gene (subinterval VIII) were the most significant contributors to the differences in the distribution of breakpoints across the a1–sh2 interval.
Statistical analysis of rates of recombination among genetic backgrounds
Recombination hot and cold spots are mostly conserved among genetic backgrounds:
As defined by Yao and Schnable (2005) "global" recombination hot and cold spots are regions that exhibit rates of recombination that are either higher or lower, respectively, than the genomewide average rate of recombination (2.4 cM/Mb). "Local" recombination hot and cold spots are regions that have rates of recombination that are higher and lower, respectively, than the average rate of recombination across the entire a1–sh2 interval in the genetic background being examined. In each of the genetic backgrounds examined in this study the average recombination rate across the a1–sh2 interval (i.e., the local average) is lower than the genomewide average rate of recombination. "Average" spots exhibit rates of recombination that do not differ significantly from the genomewide average.
All three recombination hot spots previously identified by Yao et al. (2002) in the same A1-LC Sh2/a1∷rdt sh2 heterozygote used in this study but in another genetic background [the a1 gene (subinterval II), the yz1 gene (subinterval VIII), and the apparently nongenic interloop (subinterval VI)] are both global and local recombination hot spots in all three genetic backgrounds (Figure 1B; Table 2). Similarly, two intergenic regions and one gene (x1) identified by Yao et al. (2002) as recombination cold spots (subintervals IX, X, and XI–XIII, Figure 1B; Table 2), are also cold spots in all three genetic backgrounds analyzed in this study.
Trans-acting factors affect rates of recombination within hot spots:
As discussed above, the a1 (subinterval II) and yz1 (subinterval VIII) genes and the nongenic interloop (subinterval VI) are conserved global recombination hot spots in each of the three genetic backgrounds (Table 2). If trans-acting modifiers affect each subinterval equally, the rates of recombination in each subinterval should be highest and lowest in the genetic backgrounds that have the largest (i.e., A632/LC) and smallest genetic distances (i.e., Oh43/LC), respectively, across the entire a1–sh2 interval (Table 1). Because this is often not true (Figure 1; Table 3), we conclude that trans-acting factor(s) that are polymorphic among the genetic backgrounds are differentially affecting rates of recombination within the conserved hot spots.
The most notable example of differential control of recombination among genetic backgrounds occurs in yz1 (subinterval VIII; Figure 1A). Among the three backgrounds, the rate of recombination across a1–sh2 is lowest in Oh43/LC (0.607 cM/Mb) and consistent with this low rate most subintervals show the lowest rate of recombination in this background (Figure 1B). However, more than half the recombination events isolated in the Oh43/LC background resolve in yz1 (Figure 1, C and D) and the corresponding rate of recombination is 23-fold higher than the rate across a1–sh2 (P-value ∼0; Table 2) in this background, making yz1 the most recombinationally active region in Oh43/LC. In contrast, only ∼40% of the breakpoints associated with recombinants isolated in the A632/LC and W64A/LC genetic backgrounds resolve within yz1 (Figure 1C). The recombination rate in yz1 is approximately seven-, approximately six-, and approximately fivefold hotter than the genomewide rate (Table 2) in the A632/LC, Oh43/LC, and W64A/LC genetic backgrounds, respectively, with no significant differences in recombination rates among the backgrounds (Figure 1B; Table 3). In each of the backgrounds, the rate of recombination in each of the subintervals of yz1 did not differ significantly from the rate of recombination for the entire gene (P-value > 0.05) and, further, the recombination rates in each of the subintervals of yz1 did not differ significantly among the backgrounds (data not shown).
Trans-acting modifiers can convert a region with an average recombination rate into a hot spot:
Although their levels of recombination activity vary, recombination hot and cold spots are generally conserved among the three genetic backgrounds. The exception, however, is subinterval VII, a nongenic region proximal to yz1. While no recombination was observed in this interval in another genetic background (Yao et al. 2002), in the A632/LC genetic background subinterval VII is approximately threefold more recombinationally active than the genomewide average (7.4 cM/Mb; P-value = 0.018; Table 2; Figure 1B) and approximately sevenfold hotter than the average rate across a1–sh2 in the A632/LC background (P-value = 7.1e−06; Table 2). In the W64A/LC background, however, subinterval VII is an average spot when compared to the genomewide average, but locally it is approximately three times more recombinationally active than the average rate across a1–sh2 (P-value = 0.03; Table 2). In Oh43/LC, subinterval VII is an average spot both globally and locally (Figure 1B; Table 2). In summary, this interval is more recombinationally active in each of the genetic backgrounds in this study compared to the sequence-identical interval in the line C background (Yao et al. 2002).
Genetic distances of chromosomal intervals on chromosome 1:
The genetic distances between a1 and sh2 differ significantly among the three genetic backgrounds tested in this study. To determine whether trans-acting modifiers that affect recombination across the a1–sh2 interval of chromosome 3 are global or region-specific modifiers of recombination, recombination was assayed in ∼700 progeny of cross 3 in two intervals on chromosome 1 (intervals 1S.1 and 1S.2) in the three genetic backgrounds. Intervals 1S.1 and 1S.2 are separated by 30.4 cM on chromosome 1S in the maize IBM mapping population (IBM_IDP+MMP_body map version 4.0; http://magi.plantgenomics.iastate.edu/cgi-bin/cmap). In both intervals, the genetic distances were highest in the A632/LC background consistent with results in a1–sh2 and, unlike in the a1–sh2 interval, lowest in W64A/LC (Table 4). Therefore, the relationships between genetic distances among the genetic backgrounds differ between the intervals examined on chromosome 1S and the a1–sh2 interval on 3L.
Genetic distances across two genetic intervals unlinked to a1–sh2 in three genetic backgrounds
Trans-acting genetic modifiers affect rates of recombination in the a1–sh2 interval:
The heterogeneity in rates of recombination observed in previous studies among maize genomes (Beavis and Grant 1991; Tulsieram et al. 1992; Fatmi et al. 1993; Williams et al. 1995) could be due to cis- and/or trans-acting genetic modifiers. In contrast, the genetic stocks used in this study ensured that elements that affect recombination in cis are the same in each background. This is, therefore, the first molecular analysis in plants, animals, or yeast for which differences in recombination across a multigenic interval can be directly attributed to trans-acting modifiers that differ among genetic backgrounds.
To enhance our ability to identify inbred backgrounds that exhibit polymorphic trans-acting factors, recombination was assayed in three genetically distinct maize inbreds. A632 is a Stiff Stalk (SS) Reid Yellow Dent inbred, while the Non-Stiff Stalk (NSS) inbreds Oh43 and W64A (http://www.maizegenetics.net/index.php?page=germplasm/lines.html) are members of the Lancaster Sure Crop and the Hy:T8:Wf9 subgrouping (Liu et al. 2003), respectively. A632 and Oh43 are among the six inbreds that contribute to ∼70% of all hybrids in the United States (Nass and Paterniani 2000).
Rates of recombination between a1 and sh2 varied up to approximately twofold among these three genetic backgrounds (Table 1). This establishes that trans-acting modifiers can affect rates of recombination and, not surprisingly, the effects of these factors vary among the inbreds within different heterotic groups.
Trans effects on a multigenic interval:
Unlike previous studies of trans-acting effects that used large genetic (and physical) intervals with uncharacterized molecular structures (Timmermans et al. 1997) or studies that compare genetic sizes among different genetic maps (Beavis and Grant 1991; Tulsieram et al. 1992; Fatmi et al. 1993; Williams et al. 1995), analysis of the well-characterized multigenic a1–sh2 interval (Figure 1A) allowed for observation of the effects of trans-acting modifiers in defined genic and intergenic regions.
Although recombination hot and cold spots are largely conserved among the genetic backgrounds (Figure 1B; Table 2), the rates within recombination hot spots are affected by trans-acting factors that are polymorphic among the backgrounds (Table 3). For example, in the Oh43/LC genetic background trans-acting factor(s) have seemingly redirected recombination events that might normally resolve within the nongenic interloop (subinterval VI) to yz1 (subinterval VIII) such that the resulting rate of recombination in yz1 does not differ significantly from the rates in the A632/LC and W64A/LC genetic backgrounds. In each of the conserved recombination hot spots, the relationships between recombination rates among the backgrounds vary (Table 3). This demonstrates that the trans-acting modifier(s) act by increasing or decreasing the rates of recombination within particular hot spots.
Trans-acting modifier(s) affecting recombination in this study are affecting not only the overall rate of recombination between a1 and sh2 but also the distributions of recombinants across this interval. This observation is in direct contrast to the trans-acting autonomous transposon MuDR that increased the rate of recombination at an a1 allele containing a Mu1 insertion but did not change the distribution of recombinant breakpoints compared to patterns observed in other a1 alleles (Yandeau-Nelson et al. 2005). This suggests, not surprisingly, that different types of trans-acting modifiers will affect recombination in different ways.
Trans-acting modifier(s) can act as hot and cold switches in a region of lower sequence similarity:
Although there are exceptions (Yao et al. 2002), in general plant genes are recombination hot spots and nongenic regions are cold spots (reviewed in Puchta and Hohn 1996; Schnable et al. 1998). Indeed, nongenic subintervals IX and XI–XIII are both local and global cold spots (Figure 1B; Table 2) in each of the three genetic backgrounds. In striking contrast, the nongenic subinterval VII is both a global and local hot spot in A632/LC, providing a second example of an apparently nongenic recombination hot spot. Because subinterval VII is only a local hot spot in the W64A/LC and an average spot in the Oh43/LC genetic backgrounds (Table 2), trans-acting modifiers not only are able to alter rates of recombination within hot spots but also can switch hot spots on and off.
Recombination hot spots have been shown to be "dialed down" by sequence heterology (cis-acting elements) in many organisms, including within the maize a1–sh2 interval (Yao and Schnable 2005), by a mechanism involving mismatch repair proteins (reviewed in Modrich and Lahue 1996; Borts et al. 2000; Evans and Alani 2000; Schofield and Hsieh 2003). At the maize bz1 locus, as little as 1.5% sequence divergence reduces rates of recombination twofold (Dooner 2002). In subinterval VII, the level of sequence heterology between the line C haplotype and the a1∷rdt-sh2 haplotype present in each genetic background is 3.1%. Even with this relatively high degree of heterology, the rate of recombination in A632 is approximately threefold higher than the average rate of recombination in the genome (7.4 vs. 2.4 cM/Mb; Table 2). This demonstrates that trans-acting modifiers in the A632/LC and W64A/LC (where recombination in this subinterval is also increased but to a lesser extent) backgrounds can overcome the suppression of recombination that often occurs between heterologous sequences. Further, this establishes that high sequence identity, a cis-acting factor, is not the only criterion that dictates whether a region is a recombination hot or cold spot.
Comparison of cis- and trans-acting effects on recombination:
Both cis- and trans-acting factors can potentially affect the rate of recombination in a given interval. Only by studying each type of factor in the absence of the other (e.g., trans in the absence of polymorphic cis effects) is it possible to assess how these two categories similarly and differently affect recombination. To that end, the a1–sh2 interval in maize is the first region for which the effects of trans-acting modifiers on recombination (this study) can be compared to the effects of cis-acting elements on recombination that were characterized among different a1–sh2 teosinte intervals introgressed into maize (Yao and Schnable 2005).
Both the cis- and trans-acting modifiers in the two studies affect recombination similarly in that in both studies most of the recombination breakpoints resolve in the proximal ∼10% of the a1–sh2 interval. Together these studies suggest that although cis and trans effects on recombination in the a1–sh2 interval are, in general, similar, the cis elements represented in the teosinte a1–sh2 intervals affect recombination to a greater extent than do the trans-acting modifiers that are polymorphic in the A632/LC, Oh43/LC, and W64A/LC genetic backgrounds. For example, in a subinterval containing the interloop (subinterval VI in this study), which is a recombination hot spot in each of the trans stocks, cis elements act to make this region an average, cold, and hot spot in the three different intervals studied. Such comparisons are limited, however, because the trans-acting modifiers in the cis studies most likely differ from those in the inbred backgrounds in this study.
Do the trans-acting factors act globally or on specific genetic intervals?
Trans-acting modifiers of meiotic recombination (reviewed in Wahls 1998) have been divided into two categories: those that affect recombination across the entire genome (i.e., global modifiers) and those that act only on specific (i.e., region-specific modifiers) genetic intervals (Catcheside 1977). Global trans-acting modifiers are most likely proteins intimately involved in the mechanisms of recombination (reviewed in van den Bosch et al. 2002). For example, in yeast rates of meiotic CO are globally reduced in mutants of DNA polymerase δ (Maloisel et al. 2004). In maize, desynaptic (Ji et al. 1999), which is involved in crossover control, affects recombination across the genome (Bass et al. 2003).
To test whether or not the trans-acting factors that affect the a1–sh2 interval are global modifiers, in each genetic background recombination was assayed in two intervals genetically unlinked to a1–sh2. In both intervals on chromosome 1, the relationships among genetic distances were the same. However, these relationships differed from that seen in the a1–sh2 interval (compare Tables 1 and 4). This could be due to the same trans-acting modifier(s) within a genetic background differentially affecting recombination in chromosomes 1S and 3L. Alternatively, different region-specific modifiers could be acting on at least portions of each of the two chromosomes. In addition, because the 1S.1 and 1S.2 intervals are most likely not sequence-identical among the backgrounds we cannot rule out the possibility that a combination of both trans- and cis-acting elements is affecting recombination in these intervals. Hence, these results suggest, but do not prove, that the trans-acting modifiers detected in this study do not have global effects and are instead region specific.
Region-specific trans-acting modifiers have been characterized in Schizosaccharomyces pombe (De Veaux et al. 1992; De Veaux and Smith 1994; Li et al. 1997; Krawchuk et al. 1999; Pryce et al. 2005) and Saccharomyces cerevisiae (Rockmill and Roeder 1990). The effects of chromatin organization on DSB formation and subsequent meiotic recombination repair have been extensively demonstrated in many organisms (reviewed in Lichten 2001; Petes 2001), including plant species (reviewed in Schuermann et al. 2005). In S. pombe, rec10, a protein involved in the formation of lateral elements, is required for the activation of some but not all M26-containing recombination hot spots (Pryce et al. 2005). In this case, higher-order chromatin structure affects recombination at a known hot spot. Trans-acting modifiers involved in chromatin organization or remodeling might be responsible for the transformation of the nongenic subinterval VII from an average spot in Oh43/LC to a hot spot in the A632/LC genetic background by opening the chromatin and allowing access by the recombination machinery. If so, trans-acting modifiers of chromatin most likely act in context of local structures (e.g., sequence motifs). Alternatively, genotype-specific patterns of crossover interference (reviewed in van Veen and Hawley 2003; Hillers 2004; Copenhaver 2005) could differently affect the distributions of recombination hot spots across a chromosome among genetic backgrounds.
Region-specific trans-acting modifiers could also include proteins (e.g., transcription factors) for which binding to specific cis-acting elements [e.g., unique sequence motifs (Wahls and Smith 1994; Kon et al. 1997; Pryce et al. 2005) or promoters] is necessary for activation of recombination hot spots as seen in α-hot spots in yeast (reviewed in Lichten and Goldman 1995; Petes 2001). Such interactions between cis-acting elements and trans-acting modifiers, if they occur in a1–sh2, must be due to differences among the trans-acting modifiers (e.g., binding specificity or affinity for the cis element) because the a1–sh2 intervals are sequence-identical among the genetic backgrounds.
We thank Dan Nettleton (Iowa State University) for advice regarding statistical analyses, Heather Smith for the isolation and purification of recombinant alleles, undergraduate students Brian Reinertson, Timothy Dunham, John Tenhunfeld, and Lakeysha Toomer for technical assistance, and David Glover (Purdue University) for the gift of the trans stocks. This research was supported in part by the National Research Initiative of the United States Department of Agriculture Cooperative State Research, Education and Extension Service, grant numbers 9701407 and 9901579 to P.S.S. and B.J.N. and 0101869 and 0300940 to P.S.S., and supported by Hatch Act and State of Iowa funds.
↵1 Present address: Pennsylvania State University, University Park, PA 16802.
Communicating editor: A. H. Paterson
Received August 8, 2005.
Accepted June 26, 2006.
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Јасмина Грковић Мејџор (Станишић, 1959) је српски академ и универзитетски професор. Важи за једног од најбољих познавалаца српског језика.

Образовање
Рођена је у Станишићу. Основну и средњу школу завршила је у Новом Саду. Била је ђак генерације друштвено-језичког смера гимназије "Јован Јовановић Змај" (1977). Дипломирала је на Катедри за јужнословенске језике Филозофског факултета у Новом Саду 1981, магистрирала палеославистику 1984. и докторирала 1990. године (наслов докторске тезе: Језик Псалтира из штампарије Црнојевића), на истом факултету (област палеославистике). У току школовања боравила је повремено у иностранству ради усавршавања знања језикâ (Бордо, Берлин, Лион, Минхен, Москва, Санкт Петербург, Кијев, Париз).

Наука
Главни научни доприноси академкиње Грковић Мејџор укључују проучавање језика српске средњовековне писмености, истраживања синтаксе и лексике старословенског језика, проучавање диглосије у српском средњем веку и функционалне раслојености књижевног језика тога доба и етимолошка и семантичка проучавања српског средњег века. Објавила је преко 20 радова у земљи и иностранству.

Од 1992. године је запослена на Одсеку за српски језик и лингвистику Филозофског факултета у Новом Саду. Изабрана је у звање доцента 1991. године, у звање ванредног професора 1995. године, а у звање редовног професора за област српски језик и лингвистика 2000. године. Од 2005. године је гостујући професор на Филозофском факултету Универзитета у Бањој Луци. Ментор је у изради низа дипломских и магистарских радова, као и 6 докторских дисертација, и члан комисија за оцену и одбрану магистарских и докторских теза из палеославистике, англистике, романистике.

За редовног члана САНУ је изабрана 5. новембра 2015 са приступном академском беседом Обрасци заклињања у историјској перспективи: лингвистички и когнитивни увиди. Председник је Старословенског одбора САНУ, представник је САНУ у Националном савету за научни и технолошки развој и у Стручно-научном већу Центра за научноистраживачки рад САНУ и Универзитета у Крагујевцу, као и члан седам одбора у склопу САНУ. Члан је Друштва за античке студије Србије и Европског лингвистичког друштва (-{Societas Linguistica Europaea}-). У Редакцији је часописа Зборник Матице српске за филологију и лингвистику, главни уредник часописа Прилози проучавању језика, уредник едиције Лингвистичке свеске, члан Стручне редакције (област филологија / лингвистика) за израду Српске енциклопедије. Била је гост на више страних универзитета, укључујући и Универзитет Харвард (на одељењу за словенске језике и литературу), Охајо Стејт Универзитету (Центар за средњовековне словенске студије), Институт за славистику Универзитета у Бечу и Инстутут за славистику Универзитета у Регензбургу.

Избор из библиографије

Референце

Спољашње везе
 Биографија на сајту САНУ
 Закон о родној равноправности угрожава уставне слободе - интервју ("Политика", 7. јун 2021)

Рођени 1959.
Српски професори
Бивши студенти Универзитета у Новом Саду
Академици САНУ

redpajama_set_name: RedPajamaWikipedia


text:
Sports and Politics / Activism -
Related categories: Health & Wellbeing / Changing the World /
Related booklists: Activism & Resistance, Women & Sport, Sports, Football, Hillsborough Disaster, The Olympics,
Soccer Vs. The State: Tackling Football and Radical Politics - by Gabriel Kuhn
Soccer has turned into a multi-billion-dollar industry. Professionalism and commercialization dominate its global image. Yet the game retains a rebellious side, maybe more so than any other sport co-opted by moneymakers and corrupt politicians. From ... more
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Soccer Vs. The State: Tackling Football and Radical Politics
by Gabriel Kuhn
The Fall of the House of Fifa - by David Conn
The Fall of the House of Fifa is the definitive story of Fifa's rise - and the most spectacular fall sport has ever seen. 1904: 'No person should be allowed to arrange matches for personal profit' - Fifa congress A century later: 'F ... more
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The Fall of the House of Fifa
by David Conn
And the Sun Shines Now: How Hillsborough and the Premier League Changed Britain - by Adrian Tempany
On 15 April 1989, 96 people were fatally injured on a football terrace at an FA Cup semi-final in Sheffield. The Hillsborough disaster was broadcast live on the BBC; it left millions of people traumatised, and English football in ruins. And the ... more
Order And the Sun Shines Now: How Hillsborough and the Premier League Changed Britain
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Hillsborough Voices: The Real Story Told by the People Themselves - by Kevin Sampson, in association with the Hillsborough Justice Campaign
On 15 April 1989 the world witnessed one of the worst football disasters occur at the Hillsborough Stadium in Sheffield: 96 people were crushed to death and another 766 injured in what later was admitted to be caused by a shocking failure by police. ... more
Order Hillsborough Voices: The Real Story Told by the People Themselves
Hillsborough Voices: The Real Story Told by the People Themselves
by Kevin Sampson, in association with the Hillsborough Justice Campaign
61 Minutes in Munich: The Story of Liverpool FC's First Black Footballer - by Howard Gayle
In April 1981, Howard Gayle was summoned from the substitutes' bench and sent on to play for Liverpool in the second leg of a European Cup semi-final at German champions Bayern Munich. The previous October, by filling the same role at Manc ... more
Order 61 Minutes in Munich: The Story of Liverpool FC's First Black Footballer
61 Minutes in Munich: The Story of Liverpool FC's First Black Footballer
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The Lost Boys: Inside Football's Slave Trade - by Ed Hawkins
From South America and Africa, kids as young as 13 are leaving poverty-stricken families for a new life in Europe, having been sold the vision of untold riches and the trappings of professional football. This is football's slave trade – the beautif ... more
Order The Lost Boys: Inside Football's Slave Trade
The Lost Boys: Inside Football's Slave Trade
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Hope: My Life in Football - by Hope Powell
Hope Powell was a skinny little black girl from South London who only ever wanted to play football. Brought up in a violent home, she was actively discouraged from playing. The original Bend It Like Beckham girl, this is the amazing story of how a ki ... more
Order Hope: My Life in Football
Hope: My Life in Football
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Brazil's Dance with the Devil (Updated Olympics Edition) - by Dave Zirin
The people of Brazil celebrated when they learned that in the space of two years their country would host the world's two largest sporting events: the World Cup in 2014 and the Olympics in 2016. Now they are protesting in numbers the country hasn't s ... more
Order Brazil's Dance with the Devil (Updated Olympics Edition)
Brazil's Dance with the Devil (Updated Olympics Edition)
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An Olympic Dream: The Story of Samia Yusuf Omar - by Reinhard Kleist
The remarkable story of Somali Olympian Samia Yusuf Omar. In 2008, 17-year-old Samia Yusuf Omar stood alongside some of the fastest women in the world on the start line of the Olympic 200m. Four years later, she boarded a refugee boat to Eur ... more
Order An Olympic Dream: The Story of Samia Yusuf Omar
An Olympic Dream: The Story of Samia Yusuf Omar
by Reinhard Kleist
Hillsborough: The Truth (Updated edition) - by Phil Scraton
This is the definitive, unique account of the disaster in which 96 men, women and children were killed, hundreds injured and thousands traumatised. It details the appalling treatment endured by the bereaved and survivors in the immediate aftermath, t ... more
Order Hillsborough: The Truth (Updated edition)
Hillsborough: The Truth (Updated edition)
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The Hillsborough Disaster: In Their Own Words - by Mike Nicholson
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Order The Hillsborough Disaster: In Their Own Words
The Hillsborough Disaster: In Their Own Words
by Mike Nicholson
The Dirty Game: Uncovering the Scandal at FIFA - by Andrew Jennings
In The Dirty Game, investigative reporter and BBC Panorama presenter Andrew Jennings, who has been heralded around the world for his decade-long pursuit of this story, uncovers the eye-watering level of fraud and criminal activity at the heart of FIF ... more
Order The Dirty Game: Uncovering the Scandal at FIFA
The Dirty Game: Uncovering the Scandal at FIFA
by Andrew Jennings
Power Games: A Political History of the Olympics - by Jules Boykoff, with a Foreword by Dave Zirin
A timely, no-holds barred, critical political history of the modern Olympic Games.� The Olympics have a checkered, sometimes scandalous, political history. Jules Boykoff, a former US Olympic team member, takes readers from the event�s nineteenth-cen ... more
Order Power Games: A Political History of the Olympics
Power Games: A Political History of the Olympics
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The Ugly Game: The Qatari Plot to Buy the World Cup - by Heidi Blake and Jonathan Calvert
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Playing as If the World Mattered: An Illustrated History of Activism in Sports - by Gabriel Kuhn
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Order Playing as If the World Mattered: An Illustrated History of Activism in Sports
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An homage to the classic Our Bodies, Ourselves, this encyclopedic, crowd-sourced compilation of essays, resources, information, and advice about the intersection of gender and bicycling covers a lot of ground�bold meditations on body parts, stories a ... more
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Our Bodies, Our Bikes
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Beating Berlusconi! - by John Graham Davies
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The Three Degrees: The Men Who Changed British Football Forever - by Paul Rees
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Shocking Brazil: Six Games That Shook the World Cup
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FC Sankt Pauli � the football club in the red-light district of Hamburg; a transvestite chairman; terraces populated by punks, pimps and prostitutes; a club run by anarchists, united under the skull and crossbones flag. This is the clich� that has be ... more
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Pirates, Punks & Politics FC St. Pauli: Falling in Love with a Radical Football Club
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Barbaric Sport: A Global Plague
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Ghost Milk: Calling Time on the Grand Project - by Iain Sinclair
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Wheels of Change: How Women Rode the Bicycle to Freedom
by Sue Macy
Outcasts United: An American Town, A Refugee Team, and One Woman's Quest to Make a Difference - by Warren St John
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More Than Just a Game: Football v Apartheid - The Most Important Football Story Ever Told
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Football Against the Enemy
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The Alliance française - a global network
The Alliance française Bruxelles-Europe forms part of the first international cultural network, namely the Alliance française, which was founded in Paris in 1883.
The network, which is coordinated by the Alliance française Foundation, consists of around 800 Alliance branches in 135 countries on five continents. Every day, their 8,000 teachers and 550,000 students keep the French language alive around the world and add to its international profile. The network's many cultural events attract audiences in excess of 2 million people each year. The Alliances françaises owe their autonomy to the fact that they are self-funded to the tune of 83%, are classed as associations operating under local law, and are supported by 8,500 volunteers.
The Alliance française Foundation
Founded in July 2007, the Alliance française Foundation holds charitable status. It ensures respect for statutes and the "Alliance française" label, a global brand recognised in more than 150 countries around the world on account of the quality of its language tuition and its role in supporting intercultural dialogue and promoting the ideas and values of La Francophonie.
It is also responsible for coordinating the global network, lending its expertise, advice and support to specific projects. The Foundation also appoints the executive directors who represent it in different countries or regions, cooperates with the Ministry of Foreign Affairs in appointing expatriate staff for the Alliances, and organises teaching and management courses as part of its continued professional development plan.
Learn more about the Alliance française network thanks to this infographics!
The Alliance française in Belgium
In his capacity as executive officer, the director of the Alliance française Bruxelles-Europe is the Alliance française Foundation's representative in Belgium. He coordinates and supports the educational, cultural and developmental activities of all the Alliances françaises in the country. The Alliance française network in Belgium is made up of the following bodies:
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We are members of EUNIC Brussels, a network of around twenty European Union National Institutes for Culture and embassy cultural services.
With the support of their networks around Europe, the members work together to plan and deliver multilateral cultural projects. Their aim is to promote European values, contribute to cultural diversity and boost cultural dialogue, interaction and lasting cooperation between cultures.
With its network of member institutes and its 15 languages, EUNIC Brussels is a fine example of European cultural cooperation. The cluster meets on a regular basis and the presidency rotates annually. Mindful of the cultural challenges of the modern world, EUNIC Brussels focuses its activities on intercultural dialogue, multilingualism and European citizenship.
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Steelers vs. Colts: 2016 Week 12 - News, Headlines and Updates
2016 Pittsburgh Steelers remain a work-in-progress
Against Dallas and Cleveland, the Steelers gave strong efforts despite not yet hitting on all cylinders.
By Bill Dundas Nov 24, 2016, 12:00pm EST
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Scott R. Galvin-USA TODAY Sports
While their 1-1 record for the past two games might not suggest it, the Pittsburgh Steelers appear to be slowly shaking off the malaise that hounded them throughout a majority of their recently snapped four-game losing streak. Despite allowing the Cowboys to come back and beat them at Heinz Field late in the fourth quarter, there was no shame in losing to a team currently performing at the top of its game. And the improved effort we saw against Dallas carried through last weekend in Cleveland, although the winless Browns certainly lack anything approaching the Cowboys' level of offensive prowess. But in both cases, the effort definitely was there, regardless of the outcomes.
If the Steelers truly are intent on reversing their inconsistency during the season's first half, tonight's game in Indy will be Step 2 in what Steelers Nation hopes is a 7-step program to drive into the playoffs. With Andrew Luck having been officially ruled out of the game, this appears to present a golden opportunity for the Steelers to grab a win which otherwise might have been a much more challenging proposition. The duel in Indy also gives the Black and Gold another week to work out the ongoing transition in personnel which it will need to carry it into the postseason and, hopefully, beyond.
We've seen some positive signs lately in terms of young talent developing in the secondary and a sudden, surprising uptick in defensive pressure on the quarterback. Considering the huge and unexpected absence of Cameron Heyward for the remainder of the 2016 campaign, extensive platooning of several defensive players in recent games appears to be creating some opportunities for Mike Tomlin and Company to better evaluate the talent they've got. Odd though it might seem, the Steelers' defense seems to finally be undergoing a transition which many expected would take place in the preseason rather than during the second half of the regular season.
In particular, Ryan Shazier and Stephon Tuitt appear to be coming into their own as NFL defensive stars. Meanwhile, Jarvis Jones is playing better football than many thought he could, while Sean Davis and Artie Burns are shoring up a much-maligned Steelers secondary, even as they continue to learn the NFL ropes under fire. Even Daniel McCullers made more than his usual cameo appearance in Cleveland. If McCullers can elevate his "nasty" to a level matching his size, Pittsburgh might have found the run stopper they formerly had in the legendary Big Snack. But regardless of how it all pans out, it's obvious there's currently some serious competition under way within this unit for the top of the depth chart. By the end of this season, we'll have a much better idea of where the Steelers' defense stands, which players have earned the right to hang around and what additional needs must be addressed in the 2017 NFL Draft.
That being stated, nobody watching the Steelers this season truly expects defense to carry this team to the postseason. As usual, Pittsburgh's ultimate fate will be carried on the broad shoulders of Ben Roethlisberger. For the most part, Ben still hasn't played the kind of football that many anticipated back in early September. And considering the nature of issues this team has had defensively, No. 7 needs to string together several solid performances between Thanksgiving and the end of the season. Obviously, Ben also needs to stay healthy (knock on wood). Otherwise, any opportunity the Steelers might have in the postseason goes up in smoke.
So the test resumes tonight against an Indianapolis Colts team minus its offensive leader. Like the must-win game in Cleveland last Sunday, this is another opportunity which the Steelers must seize to continue building toward a postseason berth. This is especially true in anticipation of their Week 13 matchup against the improved New York Giants at Heinz Field. What Steelers Nation truly hopes to witness tonight beneath the dome at Lucas Oil Stadium is another inspired performance on both sides of the ball, regardless of who suits up for Indy.


The Kick-Off Meeting for the ERA-NET Plus Infravation 2014 Call projects was held on 12-13th November 2015. The meeting included dynamic keynote speakers from the Infravation Steering Group and the European Commission who outlined the background and context to the Call, as well as an introduction by project coordinators of each of the selected projects per challenge. More details will follow shortly but the presentations from both days can already be accessed here.


Been using for several days and everything works well with a Hitachi Travelstar 320GB on my Dell E6400. Installation was easy. Remember to Initialize the drive so that Windows will recognize it (go to Windows disk management tools to initialize the drive as a NTFS disk). I received the caddy very quickly (3 days on the East Coast using the basic shipping option).


"The Boy With The Hammer In The Paper Bag" is the first song on Mark Eitzel's 7th album, 2001's The Invisible Man.
stands up to cheer, then you stand up to cheer.
it's the best look that we've seen all year.
to see a hunger for a pain that's endless."
too fucked up to feel anything.
If that's what you want - well, it's nothing.
no one's gonna catch you if you fall all alone.
But if that's what you want, it's nothing.
This page was last edited on 23 March 2017, at 00:34.