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Sex-chromosome dosage compensation

Dosage compensation is the process by which organisms equalize the expression of genes between members of different biological sexes. Across species, different sexes are often characterized by different types and numbers of sex chromosomes. In order to neutralize the large difference in gene dosage produced by differing numbers of sex chromosomes among the sexes, various evolutionary branches have acquired various methods to equalize gene expression among the sexes. Because sex chromosomes   contain different numbers of genes, different species of organisms have developed different mechanisms to cope with this inequality. Replicating the actual gene is impossible; thus organisms instead equalize the expression from each gene. For example, in humans, female (XX) cells randomly silence the transcription of one X chromosome, and transcribe all information from the other, expressed X chromosome. Thus, human females have the same number of expressed X-linked genes per cell as do human males (XY), both sexes having essentially one X chromosome per cell, from which to transcribe and express genes.

Different lineages have evolved different mechanisms to cope with the differences in gene copy numbers between the sexes that are observed on sex chromosomes. Some lineages have evolved dosage compensation, an epigenetic mechanism which restores expression of X or Z specific genes in the heterogametic sex to the same levels observed in the ancestor prior to the evolution of the sex chromosome. Other lineages equalize the expression of the X- or Z- specific genes between the sexes, but not to the ancestral levels, i.e. they possess incomplete compensation with "dosage balance". One example of this is X-inactivation which occurs in humans. The third documented type of gene dose regulatory mechanism is incomplete compensation without balance (sometimes referred to as incomplete or partial dosage compensation). In this system gene expression of sex-specific loci is reduced in the heterogametic sex i.e. the females in ZZ/ZW systems and males in XX/XY systems.

There are three main mechanisms of achieving dosage compensation which are widely documented in the literature and which are common to most species. These include random inactivation of one female X chromosome (as observed in humans and Mus musculus; this is called X-inactivation), a two-fold increase in the transcription of a single male X chromosome (as observed in Drosophila melanogaster), and decreased transcription by half in both of the X chromosomes of a hermaphroditic organism (as observed in Caenorhabditis elegans). These mechanisms have been widely studied and manipulated in model organisms commonly used in the laboratory research setting. A summary of these forms of dosage compensation is illustrated below. However, there are also other less common forms of dosage compensation, which are not as widely researched and are sometimes specific to only one species (as observed in certain bird and monotreme species).

Random inactivation of one ♀ X
One logical way to equalize gene expression amongst males and females that follow a XX/XY sex differentiation scheme would be to decrease or altogether eliminate the expression of one of the X chromosomes in an XX, or female, homogametic individual, such that both males and females then express only one X chromosome. This is the case in many mammalian organisms, including humans and mice.

The evidence for this mechanism of dosage compensation was discovered prior to scientists' understanding of what its implications were. In 1949, Murray Barr and Ewert Bertram published data describing the presence of "nucleolar satellites, which they observed were present in the mature somatic tissue of different female species. Further characterization of these satellites revealed that they were actually packages of condensed heterochromatin, but a decade would pass before scientists grasped the significance of this specialized DNA.

Then, in 1959 Susumu Ohno proved that these satellite-like structures found exclusively in female cells were actually derived from female X chromosomes. He called these structures Barr bodies after one of the investigators who originally documented their existence. Ohno's studies of Barr bodies in female mammals with multiple X chromosomes revealed that such females used Barr bodies to inactivate all but one of their X chromosomes. Thus, Ohno described the "n-1" rule to predict the number of Barr bodies in a female with n number of X chromosomes in her karyotype.

Simultaneously, Mary F. Lyon began investigating manipulations of X-linked traits that had phenotypically visible consequences, particularly in mice, whose fur color is a trait intimately linked to the X chromosome. Building on work done by Ohno and his colleagues, Lyon eventually proved that either the maternal or paternal X chromosome is randomly inactivated in every cell of the female body in the species she was studying, which explained the heterogeneous fur patterns she observed in her mosaic mice. This process is known as X-inactivation, and is sometimes referred to as "lyonization". This discovery can be easily extrapolated to explain the mixed color patterns observed in the coats of tortoiseshell cats. The fur patterns characteristic of tortoiseshell cats are found almost exclusively in females, because only they randomly inactivate one X chromosome in every somatic hair cell. Thus, presuming that hair color determining genes are X-linked, it makes sense that whether the maternal or paternal X chromosome is inactivated in a particular hair cell can result in differential fur color expression.

Compounding on Lyon's discoveries, in 1962 Ernest Beutler used female fibroblast cell lineages grown in culture to demonstrate the heritability of lyonization or random X-inactivation. By analyzing the differential expression of two existing, viable alleles for the X-linked enzyme glucose-6-phosphate dehydrogenase (G6PD) gene, Beutler observed that the inactivation of the gene was heritable across passaged generations of the cells.

This pattern of dosage compensation, caused by random X-inactivation, is regulated across development in female mammals, following concerted patterns throughout development; for example, at the beginning of most female mammal development, both X chromosomes are initially expressed, but gradually undergo epigenetic processes to eventually achieve random inactivation of one X. In germ cells, inactivated X chromosomes are then once again activated to ensure their expression in gametes produced by female mammals.

Thus, dosage compensation in mammals is largely achieved through the silencing of one of two female X chromosomes via X-inactivation. This process involves histone tail modifications, DNA methylation patterns, and reorganization of large-scale chromatin structure encoded by the X-ist gene. In spite of these extensive modifications, not all genes along the X chromosome are subject to X-inactivation; active expression at some loci is required for homologous recombination with the pseudo-autosomal region (PAR) of the Y chromosome during meiosis. Additionally, 10-25% of human X chromosome genes, and 3-7% of mouse X chromosome genes outside of the PARs show weak expression from the inactive X chromosome.

Random X-inactivation demands that the cell can determine if it contains more than one active X-chromosome before acting to silence any extraneous X-chromosome(s). This process is known as "counting". The exact molecular mechanism of counting is still unknown, but a popular model posits that autosomes produce factors that repress X-inactivation, while X-chromosome products that promote X-inactivation. These two conflicting forces are balanced such that if there is more than one X-chromosome X-inactivation will occur, but if there is only one, the autosomal products will successfully prevent the process.

Not all random X-inactivation is entirely random. Some alleles, generally mutations in the X-inactivation center on the X-chromosome have been demonstrated to confer a bias towards inactivation for the chromosome on which they sit. Truly random X-inactivation may also appear to be non-random if one X-chromosome carries a deleterious mutation. This can result in fewer cells which express the lower-fitness X-chromosome to be present in the body as these cells are selected against.

Two-fold increased transcription of a single ♂ X
Another mechanism common for achieving equal X-related genetic expression between males and females involves two-fold increased transcription of a single male X chromosome. Thus, heterogametic male organisms with one X chromosome may match the level of expression achieved in homogametic females with two active X chromosomes. This mechanism is observed in Drosophila.

The concept of dosage compensation actually originated from an understanding of organisms in which males upregulated X-linked genes two-fold, and was much later extended to account for the observation of the once mysterious Barr bodies. As early as 1932, H.J. Muller carried out a set of experiments which allowed him to track the expression of eye color in flies, which is an X-linked gene. Muller introduced a mutant gene that caused loss of pigmentation in fly eyes, and subsequently noted that males with only one copy of the mutant gene had similar pigmentation to females with two copies of the mutant gene. This led Muller to coin the phrase "dosage compensation" to describe the observed phenomenon of gene expression equalization.

Despite these advances, it was not until Ardhendu Mukherjee and W. Beermann performed more advanced autoradiography experiments in 1965 that scientists could confirm that transcription of genes in the single male X chromosome was double that observed in the two female X chromosomes. Mukherjee and Beermann confirmed this by designing a cellular autoradiography experiment that allowed them to visualize incorporation of [3H]uridine into ribonucleic acid of the X chromosomes. Their studies showed equal levels of [3H]uridine incorporation in the single male X chromosome and the two female X chromosomes. Thus, the investigators concluded that the two-fold increase in the rate of RNA synthesis in the X chromosome of the male relative to those of the female could account for Muller's hypothesized dosage compensation.

In the case of two-fold increased transcription of a single male X chromosome, there is no use for a Barr body, and the male organism must use different genetic machinery to increase the transcriptional output of their single X chromosome. It is common in such organisms for the Y chromosome to be necessary for male fertility, but not for it to play an explicit role in sex determination. In Drosophila, for example, the sex lethal (SXL) gene acts as a key regulator of sexual differentiation and maturation in somatic tissue; in XX animals, SXL is activated to repress increased transcription, while in XY animals SXL is inactive and allows male development to proceed via increased transcription of the single X. Several binding sites exist on the Drosophila X chromosome for the dosage compensation complex (DCC), a ribonucleoprotein complex; these binding sites have varying levels of affinity, presumably for varying expression of specific genes. The Male Specific Lethal complex, composed of protein and RNA binds and selectively modifies hundreds of X-linked genes, increasing their transcription to levels comparable to female D. melanogaster.

In organisms that use this method of dosage compensation, the presence of one or more X chromosomes must be detected early on in development, as failure to initiate the appropriate dosage compensation mechanisms is lethal. Male specific lethal proteins (MSLs) are a family of four proteins that bind to the X chromosome exclusively in males. The name "MSL" is used because mutations in these genes cause inability to effectively upregulate X-linked genes appropriately, and are thus lethal to males only and not their female counterparts. SXL regulates pre-messenger RNA in males to differentially splice MSLs and result in the appropriate increase in X chromosome transcription observed in male Drosophila. The immediate target of SXL is male specific lethal-2 (MSL-2). Current dogma suggests that the binding of MSL-2 at multiple sites along the SXL gene in females prevents proper MSL-2 translation, and thus, as previously stated, represses the possibility for X-linked genetic upregulation in females. However, all other transcription factors in the MSL family—maleless, MSL-1, and MSL-3—are able to act when SXL is not expressed, as in the case in males. These factors act to increase male X chromosome transcriptional activity. Histone acetylation and the consequent upregulation of X-linked genes in males is dictated by the MSL complex. Specifically, special roX non-coding RNAs on the MSL complexes facilitate binding to the single male X chromosome, and dictate acetylation of specific loci along the X chromosome as well as the formation of euchromatin. Though these RNAs bind at specific sites along the male X chromosome, their effects spread along the length of the chromosome and have the ability to influence large-scale chromatin modifications. The implications of this spreading epigenetic regulation along the male X chromosome is thought to have implications for understanding the transfer of epigenetic activity along long genomic stretches.

Decreased transcription of both hermaphroditic Xs by half
Other species that do not follow the previously discussed conventions of XX females and XY males must find alternative ways to equalize X-linked gene expression among differing sexes. For example, in Caenorhabditis elegans (or C. elegans), sex is determined by the ratio of X chromosomes relative to autosomes; worms with two X chromosomes (XX worms) develop as hermaphrodites, whereas those with only one X chromosome (XO worms) develop as males. This system of sex determination is unique, because there is no male specific chromosome, as is the case in XX/XY sex determination systems. However, as is the case with the previously discussed mechanisms of dosage compensation, failure to express X-linked genes appropriately can still be lethal.

In this XX/XO sex determination system, gene expression on the X chromosome is equalized by downregulating expression of genes on both X chromosomes of hermaphroditic XX organisms by half. In these XX organisms, the dosage compensation complex (DCC) is assembled on both X chromosomes to allow for this tightly regulated change in transcription levels. The DCC is often compared to the condensin complex, which is conserved across the mitotic and meiotic processes of many species. This complex is crucial to the condensation and segregation of chromosomes during both meiosis and mitosis. Because data substantiates the theory that dosage compensation in other species is caused by chromatin-wide modifications, many theorize that the DCC in particular functions similar to the condensin complex in its ability to condense or remodel the chromatin of the X chromosome.

The role of the DCC in this form of dosage compensation was postulated by Barbara J. Meyer in the 1980s, and its individual components and their cooperative function were later parsed out by her lab. Notably, in 1999, data from Meyer's lab showed that SDC-2 is a particularly important transcriptional factor for targeting the DCC to the X chromosome and for assembling DCC components onto the X chromosomes in XX embryos. More recently, Meyer's lab has shown that proteins known as X-linked signal elements (XSEs) operate in concert with SDC-2 to differentially repress and activate other genes in the dosage compensation pathway. By selectively mutating a panel of genes hypothesized to contribute to dosage compensation in worms, Meyer's group demonstrated which XSEs specifically play a role in determining normal dosage compensation. They found that during embryonic development, several X-linked genes—including sex-1, sex-2, fox-1, and ceh-39—act in a combinatorial fashion to selectively repress transcriptional activity of the xol-1 gene in hermaphrodites. Xol-1 expression is tightly regulated during early development, and is considered the most upstream gene in sex determination of C. elegans. In fact, xol-1 is often referred to in the literature as the master sex regulatory gene of C. elegans. XX C. elegans embryos have much lower xol-1 expression than their XO counterparts, resulting from overall increases in the amount of SEX-1, SEX-2, CEH-39, and FOX-1 transcription produced in the female embryos. This consequent decrease in xol-1 expression then allows higher SDC-2 expression levels, which aids in the formation and function of the DCC complex in the XX hermaphroditic worms, and in turn results in equalized expression of X-linked genes in the hermaphrodite.

Though all of the above-mentioned XSEs act to reduce xol-1 expression, experimentally reducing expression levels of these individual XSEs has been shown to have a minimal effect on sex determination and successful dosage compensation. This could be in part because these genes encode different proteins that act cooperatively rather than in an isolated fashion; for example, SEX-1 is a nuclear hormone receptor, while FOX-1 is an RNA-binding protein with properties capable of inducing post-transcriptional modifications in the xol-1 target. However, reducing the level of more than one XSE in different combinational permutations seems to have an additive effect on ensuring proper sex determination and resultant dosage compensation mechanics. This supports the hypothesis that these XSEs act together to achieve the desired sex determination and dosage compensation fate. Thus, in this model organism, the achieved level of X-chromosome expression is directly correlated to the activation of multiple XSEs that ultimately function to repress xol-1 expression in a developing worm embryo. A summary of this C. elegans mechanism of dosage compensation is illustrated below.

Other species-specific methods
The ZZ/ZW sex system is used by most birds, as well as some reptiles and insects. In this system the Z is the larger chromosome so the males (ZZ) must silence some genetic material to compensate for the female's (ZW) smaller W chromosome. Instead of silencing the entire chromosome as humans do, male chickens (the model ZZ organism) seem to engage in selective Z silencing, in which they silence only certain genes on the extra Z chromosome. Thus, male chickens express an average of 1.4-1.6 of the Z chromosome DNA expressed by female chickens. The Z chromosome expression of male zebra finches and chickens is higher than the autosomal expression rates, whereas X chromosome expression in female humans is equal to autosomal expression rates, illustrating clearly that both male chickens and male zebra finches practice incomplete silencing. Few other ZZ/ZW Systems have been analyzed as thoroughly as the chicken; however a recent study on silkworms revealed similar levels of unequal compensation across male Z chromosomes. Z-specific genes were over-expressed in males when compared to females, and a few genes had equal expression in both male and female Z chromosomes.

In chickens, most of the dosage compensated genes exist on the Zp, or short, arm of the chromosome while the non-compensated genes are on the Zq, or long, arm of the chromosome. The compensated (silenced) genes on Zp resemble a region on the primitive platypus sex chromosome, suggesting an ancestor to the XX/XY system.

Birds
The sex chromosomes of birds evolved separately from those of mammals and share very little sequence homology with the XY chromosomes. As such, scientists refer to bird sex chromosomes as a ZW sex-determining system, with males possessing two Z chromosomes, and females possessing one Z chromosome and one W. Thus, dosage compensation in birds could be hypothesized to follow a pattern similar to the random X-inactivation observed in most mammals. Alternatively, birds might show decreased transcription of the two Z chromosomes present in the male heterogametic sex, similar to the system observed in the two hermaphrodite X chromosomes of C. elegans. However, bird mechanisms of dosage compensation differ significantly from these precedents. Instead, male birds appear to selectively silence only a few genes along one of their Z chromosomes, rather than randomly silencing one entire Z chromosome. This type of selective silencing has led some people to label birds as "less effective" at dosage compensation than mammals. However, more recent studies have shown that those genes on the Z chromosome which are not inactivated in birds may play an important role in recruiting dosage compensation machinery to the Z chromosome in ZZ organisms. In particular, one of these genes, ScII has been demonstrated to be an ortholog of xol-1, the master sex regulator gene in C. elegans. Thus, the function of the selective silencing may be to spare dosage compensation of genes crucial for sex determination of homologous pairing.

While the epigenetic mechanisms behind dosage compensation in birds are poorly understood, especially in comparison to the well-studied mechanisms of dosage compensation in humans and Drosophila, several recent studies have revealed promising sequences. One example is MHM (male hypermethylated) RNA, an Xist-like long noncoding RNA that is expressed only in female chickens (ZW). It is associated with female-specific hyper-acetylation of lysine 16 on histone 4 near the MHM locus on the Z chromosome. This MHM locus is heavily studied as a site of dosage compensation because male Z chromosomes are hypermethylated and thus underexpress genes in this area in comparison to female Z chromosomes which are hyperacetylated and overexpress these genes. There has been debate on whether the MHM locus constitutes dosage compensation, however, since scientists claim that even if the MHM locus has been found to have significantly greater expression in females than in males, it could not even be considered to be a dosage compensation mechanism since it does not balance gene dose between the Z chromosome and autosomes in the heterogametic sex.

Similar to mammals, chickens seem to use CpG islands (segments of Cytosine-phosphate-Guanine that are more readily methylated and silenced than other DNA segments) to regulate gene expression. One study found that CpG islands were found primarily in compensated areas of the Z chromosome—areas that are differentially expressed in male and female chickens. Thus it is likely that these CpG islands are locations where genes on the male Z chromosome are methylated and silenced, but which stay functional on the female Z chromosome.

Monotremes
Monotremes are a class of basal mammals that also lay eggs. They are an order of mammals that includes platypuses and four species of echidna, all of which are egg-laying mammals. While monotremes use an XX/XY system, unlike other mammals, monotremes have more than two sex chromosomes. The male short-beaked echidna, for example, has nine sex chromosomes—5 Xs and 4 Ys, and the male platypus has 5 Xs and 5 Ys.

Platypuses are a monotreme species whose mechanism of sex determination has been extensively studied. There is some contention in academia about the evolutionary origin and the proper taxonomy of platypuses. A recent study revealed that four platypus X chromosomes, as well as a Y chromosome, are homologous to some regions on the avian Z chromosome. Specifically, platypus X1 shares homology with the chicken Z chromosome, and both share homology with the human chromosome 9. This homology is important when considering the mechanism of dosage compensation in monotremes. In 50% of female platypus cells, only one of the alleles on these X chromosomes is expressed while in the remaining 50% multiple alleles are expressed. This, combined with the portions that are homologous to chicken Z and human 9 chromosomes imply that this level of incomplete silencing may be the ancestral form of dosage compensation.

Regardless of their ambiguous evolutionary history, platypuses have been empirically determined to follow an XY sex-determination system, with females possessing five pairs of X chromosomes as the homogametic sex, and males possessing five X and five Y chromosomes as the heterogametic sex. Because the entire genome of the platypus has yet to be completely sequenced (including one of the X chromosomes), there is still continued investigation as to the definitive mechanism of dosage compensation that Platypuses follow. Research from the laboratory of Jennifer Graves used qPCR and SNP analysis of BACs containing various genes from X chromosomes in order to find whether multiple alleles for particular X-linked genes were being expressed at once, or were otherwise being dosage compensated. Her group found that in female platypuses, some X-linked genes only expressed an allele from one X chromosomes, while other genes expressed multiple alleles. This appears to be a system similar to the selective silencing method of dosage compensation observed in birds. However, about half of all X-linked genes also seemed to stochastically express only one active copy of said gene, alluding to the system of random X-inactivation observed in humans. These findings suggest that platypuses may employ a hybrid form of dosage compensation that combines feature from mammals as well as birds. Understanding the evolution of such a system may have implications for solidifying the true ancestral lineage of monotremes.

Plants
In addition to humans and flies, some plants also make use of the XX/XY dosage compensation systems. Silene latifolia plants are also either male (XY) or female (XX), with the Y chromosome being smaller, with fewer genes expressed, than the X chromosome. Two separate studies have shown male S. latifolia expression of X-linked genes to be about 70% of the expression in females. If the S. latifolia did not practice dosage compensation, the expected level of X-linked gene expression in males would be 50% that of females, thus the plant practices some degree of dosage compensation but, because male expression is not 100% that of females, it has been suggested that S. latiforia and its dosage compensation system is still evolving. Additionally, in plant species that lack dimorphic sex chromosomes, dosage compensation can occur when aberrant meiotic events or mutations result in either aneuploidy or polyploidy. Genes on the affected chromosome may be upregulated or down-regulated to compensate for the change in the normal number of chromosomes present.

Reptiles
Research into dosage compensation has been carried out in six species of toxicoferan reptiles and in one species of softshell turtle. Two species of caenophidian snake (one which belongs to the family Viperidae and the other to the family Colubridae) have been investigated and both of these exhibit female heterogametic sex determination systems (ZZ\ZW) and have incomplete compensation without balance. The Komodo dragon exhibits incomplete compensation without dosage balance in their independently evolved ZZ/ZW system. In the XX/XY system of Basiliscus vittatus and multiple neo-sex chromosomes with male heterogamety in the pygopodid gecko Lialis burtonis incomplete compensation without dosage balance were also seen. The Green anole (Anolis carolinensis; Dactyloidea), has XX/XY sex determination and unlike the other squamates studied to date has complete dosage compensation with dosage balance. In the Florida softshell turtle (Apalone ferox) with ZZ/ZW sex chromosomes, the lack of dosage balance in the expression of Z-linked genes was also found.

X chromosome inactivation and embryonic stem cells
XCI is initiated very early during female embryonic development or upon differentiation of female embryonic stem (ES) cells and results in inactivation of one X chromosome in every female somatic cell. This process is initiated very early during development, around the two- to eight-cell stage and is maintained in the developing extra-embryonic tissues of the embryo, including the fetal placenta. Xist RNA induces heterochromatinization of the X chromosome by attracting chromatin modifiers, involved in gene silencing. Xist RNA is tightly associated with the Xi and it is required for X Chromosome Inactivation to occur in cis. Knockout studies in female ES cells and mice have shown that X chromosomes bearing a deletion of the Xist gene are unable to inactivate the mutated X. Most of the human female ES cell lines display an inactivated X chromosome already in the undifferentiated state characterized by XIST expression, XIST coating and accumulated markers of heterochromatin on the Xi.

It is widely thought that human embryos do not employ XCI prior to implantation. Female embryos have an accumulation of Xist RNA on one of the two X chromosomes, beginning around the 8-cell stage. Xist RNA accumulates at the morula and blastocyst stages and is shown to be associated with transcriptional silencing of the Xist-coated chromosomal region, therefore indicating dosage compensation has occurred. Recently, however, it has become increasingly apparent that XCI of the paternal X chromosome is already present from the 4-cell stage onward in all cells of preimplantation mouse embryos, not the 8-cell stages.

Xist, Xite, and Tsix and their roles in X-inactivation
Xite and Xist, are both long non-coding RNAs that regulate and facilitate the process of X-inactivation and are important in the silencing of genes within the X chromosome that is being inactivated. These work in combination with Tsix, which is non-coding RNA that is an antisense which downregulates the effects of Xist on the X chromosome in which it is expressed on the maternal X chromosome upon the start regulation of X-inactivation. These three RNAs regulate the X-X pair in a cisorientation in order to be able to have both chromosomes available for inhibitory actions. Tsix and Xite have basic lncRNAs functions in addition to X-inactivation and regulate the X-X pair in the transorientation. This ensures exclusive silencing for both X chromosomes. Xite and Tsix are both essential within the orientational directional processes in cis and trans as it is seen that without Tsix and Xite in trans it perturbs pairing and counting of genes.

Once the Xist is turned off and no longer regulates the process, the Tsix will slowly decrease in expression as well until both RNAs are no longer being changed by Xic. Xite is the locus that harbors intergenic transcription start sites from hypersensitive sites of allelic crossovers/differences. When X-inactivation begins, the transcription of Xite increases and signals for the downregulation of Tsix in cisorientation, which is on the silent X chromosome, all while promoting the Tsix persistence on the active X chromosome. Xite also has major roles to play in the asymmetry of Tsix expression and generates X chromosome inequality through moving and helping orient the chromosomes to be acted upon by the correct subsequent lncRNA, either Tsix or Xist.

Neo-sex chromosomes and dosage compensation
The monarch butterfly Danaus plexippus belongs to the order Lepidoptera and has 30 chromosomes one of which is a neo-sex chromosome which is the result of a fusion between one of the sex chromosomes and an autosome. A study using a combination of methods (Hi-C assembly, coverage analysis and ChIp-seq) found that the neo-Z segment exhibits complete dosage compensation which is achieved by increased transcription in ZW females. Interestingly, the ancestral Z segment exhibits dosage balance with transcription levels being equal between both genders but less than the expected ancestral level, and this is achieved by decreased transcription in ZZ males.

title: Sex-chromosome dosage compensation


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Federal Mediation and Conciliation Service (United States)

The Federal Mediation and Conciliation Service (FMCS), founded in 1947, is an independent agency of the United States government, and the nation's largest public agency for dispute resolution and conflict management, providing mediation services and related conflict prevention and resolution services in the private, public, and federal sectors. FMCS is tasked with mediating labor disputes around the country; it provides training and relationship development programs for management and unions as part of its role in promoting labor-management peace and cooperation. The Agency also provides mediation, conflict prevention, and conflict management services outside the labor context for federal agencies and the programs they operate. The FMCS headquarters is located in Washington, D.C., with other offices across the country.

About FMCS
FMCS was created by Congress as a neutral and independent government agency upon enactment of the Labor-Management Relations Act of 1947 (Taft–Hartley Act) and mandated to resolve industrial conflict and promote labor-management peace and cooperation, minimizing the impact of these disputes on the free flow of commerce. With its headquarters in Washington, D.C., and offices across the country, the agency has, for decades, been providing dispute resolution and conflict management services for employers and unions across industries and work activities in the private, public, and federal sectors. FMCS has also been involved in facilitating negotiated rulemaking processes and providing conflict prevention and conflict management for a wide range of agencies in the federal sector. The Agency also has an international program, partnering with more than 60 countries to provide consulting and training in labor dispute resolution and the design of conflict management systems.

Role under the Taft–Hartley Act
The Federal Mediation and Conciliation Service was created as an independent agency of the federal government under the terms of the Labor Management Relations Act of 1947 (better known as the Taft–Hartley Act) to replace the United States Conciliation Service that previously operated within the Department of Labor. Under Taft-Hartley, FMCS may offer its services "in any labor dispute in any industry affecting commerce, either upon its own motion or upon the request of one or more of the parties to the dispute, whenever in its judgment such dispute threatens to cause a substantial interruption of commerce." By statute, FMCS receives advance notification any time a party to a collective bargaining agreement intends to terminate or modify the contract upon expiration. No modification or termination of a collective bargaining agreement is permitted unless the party wishing to modify or terminate notifies the other party at least 60 days prior to expiration and, within 30 days after notice to the other party, notifies FMCS and applicable state mediation agencies. For healthcare institutions, the notice times are extended to 90 and 60 days respectively.

FMCS also has a large labor arbitration program. The Agency maintains a roster of approximately 1,000 private arbitrators who are vetted based on their background, experience, and training in issues arising under collective bargaining agreements. Typically, more than 10,000 requests for arbitrator panels are received by FMCS each year from parties to labor-management grievance disputes. Panels are randomly drawn from the FMCS Roster based on specified parameters, and the parties select an arbitrator who is then officially appointed by FMCS. Arbitrators must abide by the Code of Professional Responsibility for Arbitrators of Labor-Management Disputes, to which FMCS is a signatory; the Code is incorporated by reference in the Agency's federal regulations.

Formation and first director
On August 7, 1947, President of the United States Harry S. Truman appointed Cyrus S. Ching as the first director of the FMCS. As Director of the FMCS, he received $12,000, placing the position at par with the National Labor Relations Board. Ching had been a member of the National War Labor Board until 1943, and had been an employee of the United States Rubber Company since 1919, serving as the firm's director of industrial and public relations in 1929. Ching would take office as of August 22, 1947, the date established in the Taft–Hartley Act for the creation of the FMCS as an independent agency, and would assume the role of the nation's top labor mediator from Edgar L. Warren, who had filled the senior mediation role for the U.S. Conciliation Service within the Labor Department. After conferring with the President in August, Ching stated that he would assume his role as director in early September upon the completion of his duties at U.S. Rubber. Ching stated that his role was to settle labor disputes at the level when and where they develop.

Ching was sworn into office on September 5, 1947, with an oath administered by Judge Henry White Edgerton at ceremonies also attended by Howard T. Colvin, who served as acting head from the August 22 creation of the FMCS, as well as other representatives of labor, industry and government.

Directors
Directors of the Federal Mediation and Conciliation Service (with the date they took office listed and the President who made the appointment shown in parentheses), are as follows:

 * 1) Cyrus S. Ching (1947; Truman)
 * 2) David L. Cole (1952; Truman)
 * 3) Whitney P. McCoy (1953; Eisenhower)
 * 4) Joseph F. Finnegan (1955; Eisenhower)
 * 5) William E. Simkin (1961; Kennedy), the longest-serving Director, departing office in 1969
 * 6) J. Curtis Counts (1970; Nixon)
 * 7) William Usery, Jr. (1973; Nixon)
 * 8) James F. Scearce (1976; Ford)
 * 9) Wayne L. Horvitz (1977; Carter)
 * 10) Kenneth Moffett (1982; Reagan), served for seven months.
 * 11) Kay McMurray (1982; Reagan)
 * 12) Bernard E. DeLury (1990; G. H. W. Bush)
 * 13) John Calhoun Wells (1993; Clinton)
 * 14) C. Richard Barnes (1999; Clinton)
 * 15) Peter J. Hurtgen (2002; G. W. Bush)
 * 16) Arthur F. Rosenfeld (2006; G. W. Bush)
 * 17) George H. Cohen (2009; Obama)
 * 18) Allison Beck (2014; Obama), the first woman to serve as director
 * 19) Richard Giacolone (2018; Donald J. Trump)

Notable roles and events
Representatives of the FMCS played a role in negotiations between Bethlehem Shipbuilding Corporation and the Marine and Shipbuilding Workers in a strike that started in June 1947.

In 1973, a Relationship-by-Objectives (RBO) program is developed for use in extreme cases of poor labor-management relations, when continued deterioration of the relationship could have drastic economic effect. The first RBO program is delivered in Maine on behalf of the Georgia-Pacific Company and Paperworkers Local 27.

In 1975, FMCS officially entered a new arena: Alternative Dispute Resolution (ADR). Congress passes Public Law 93-531, directing the Service to mediate a 100-year old land dispute between the Hopi and Navajo Indian Tribes in Arizona.

In 1978, Congress extended the FMCS charter to mediate disputes beyond the private sector to the Federal government.

In November,1979, FMCS began mediating age discrimination complaints.

In 1983, FMCS was the first agency to provide the service of negotiated rulemaking, or “reg-neg.” Conducted with the Federal Aviation Administration and Department of Transportation, regulations were developed to deal with flight and duty time of pilots.

The institutionalization of Alternative Dispute Resolution (ADR) came with the passage of the ADR Act of 1990 which created the responsibility of every federal agency to look at its mission and to see what could be resolved through ADR techniques; also to establish an ADR coordinator, and to promote these efforts. FMCS testified and described the kind of work it did both for promoting ADR and negotiated rulemaking.

In 1996, representatives from FMCS facilitated an ADR process in Minnesota regarding land use issues in the Boundary Waters Canoe Area Wilderness (BWCAW) and in Voyageurs National Park (VNP). In addition to agreements reached on the BWCAW, the mediation team also announced agreements on strategies to handle problems in the park having to do with public safety, improved Park Service consultation with local people, and other issues.

In 1997, representatives from FMCS mediated negotiations between United Parcel Service and the International Brotherhood of Teamsters. It required three weeks of mediation to bring an end to the largest national strike in two decades.

In June 2002, representatives from FMCS facilitated National Institute of Standards and Technology (NIST) discussions in New York City regarding the scope of the subsequent building and fire safety investigation following the Sept. 11, 2001 terrorist attacks. The meetings, which were conducted over the course of five sessions, featured more than 30 speakers, many of whom had family members and friends who were killed in the collapse of the World Trade Center towers. The impact of FMCS’ role was a successful discussion which provided important data for NIST's assessment and investigation.

Representatives of the FMCS mediated negotiations between the National Football League and the National Football League Players Association in contract talks in February 2011.

In November 2012, the National Hockey League and National Hockey League Players Association agreed to submit their negotiations to the FMCS in an effort to resolve the 2012 NHL lockout.

In December 2015, representatives from FMCS facilitated a regulatory negotiations process with the US Department of Energy involving industry, labor groups, and environmentalists to help produce the biggest energy savings standards in US history.

In 2015, representatives of the FMCS mediated negotiations and agreement between the International Longshore and Warehouse Union (ILWU) and the Pacific Maritime Association covering operations at 29 U.S. ports on the Pacific coast. Subsequently, an extension to the contract was announced in with a July 1, 2022 expiration date.

In February 2019, representatives of the FMCS mediated negotiations and agreement between the Denver Classroom Teachers Association and Denver Public School District, ending not only “the first teachers' strike in Denver in 25 years -- it concludes 15 months of sometimes acrimonious negotiations.”

From January to April 2019, FMCS facilitated regulatory negotiations meetings with the Department of Education. The meetings proposed changes/new regulations for the Federal Student Aid programs authorized under Title IV of the Higher Education Act of 1965 as amended. These particular negotiations were on the topic of Accreditation and Innovation, including TEACH grant requirements, Distance Learning and Faith-Based Institutions. The meetings were live-streamed and open to the public with time at the end of each day for public comment.

In April 2019, FMCS and the Equal Employment Opportunity Commission (EEOC) officially committed to work together to resolve federal workplace disputes by utilizing ADR as a means of efficiently reducing the backlog of federal sector charges,

In June 2019, FMCS and the Federal Labor Relations Authority (Authority) announced a new commitment to work together to provide labor organizations and agencies with an opportunity to resolve negotiability appeals before they are considered by the Authority's Members for a decision.

In August 2019, representatives of FMCS mediated negotiations between the State of Alaska and the Inlandboatmen's Union, ending a nine-day strike.

In August 2020, representatives from Bath Iron Works and International Association of Machinists Local S6 signed an agreement with FMCS’ assistance to end the shipyard worker strike in Bath, Maine. The 63-day strike drew national attention against the backdrop of a global pandemic and in an election year.

Shared Neutrals program
In 2019, FMCS began administrative management of the Federal Shared Neutrals Program, an interagency collaborative effort in support of alternative dispute resolution, formerly operated as Sharing Neutrals by the U.S. Department of Health and Human Services (HHS). FMCS administers the program in the National Capital Region, in coordination with participating federal agencies that contribute to, and make use of, a pool of collateral-duty federal employees to mediate cases outside their own agency. In many regions of the country, Federal Executive Boards (FEBs) have created individual programs modeled after the DC-based program but run through each FEB. These programs continue separate operations, except for those who may wish to have FMCS administer their regional Shared Neutrals program.

Annually, the Shared Neutrals Program provides workplace mediation services for hundreds of cases across more than 50 participating agencies and sub-agencies, supported by a pool of more than 250 collateral-duty federal employees who are dedicated to assisting in the resolution of workplace disputes.

Awards and accolades
FMCS received the top ranking among small Federal agencies as a Best Place to Work in the Federal Government, according to the 2018 rankings by the nonprofit Partnership for Public Service and Boston Consulting Group.

With rankings first being held in 2003, 2018 marks the fifth time FMCS was selected for the top spot, having also earned this distinction in 2005, 2007, 2015, and in 2017. Additionally, the Agency has consistently ranked in the top 4 since the “small agency” category rankings were first included in 2007. Data points from the FEVS revealed that FMCS's 2018 employee engagement score is 87.2 out of 100, which placed FMCS at 1 out of 29 small agencies. By 2022, the score had dropped to 63.6 out of 100, placing FMCS as the 24th out of 30 small agencies.

National Labor-Management Conference (NLMC)
FMCS hosts a biennial conference to promote better relationship and dispute management as a proactive means for preventing conflict that can impair organizational success. Historically, the NLMC attracts over 1000 industrial relations professionals, representatives of labor and management, academics, arbitrators and legal professionals across the labor relations and employment spectrum. The conference was most recently held virtually in 2020 due to the COVID-19 pandemic. The next conference, “Challenges and Choices at Work in a Time of Heightened Worker Activism”, is scheduled for June 26-30, 2024.

2013 Washington Examiner Series
In 2013, The Washington Examiner alleged in a series of articles that employees at the agency had made improper purchases such as auto leases and spouses' cellular phones using government credit cards.

In response, the FMCS issued the following statement: "These items which the Examiner is inquiring about appear to have been the subject of a now-settled employment dispute involving a disgruntled FMCS employee. These purchasing issues were addressed in the settlement, but must remain confidential under federal personnel rules, as noted, in the absence of a release from the employee.

"When FMCS became aware of this employee’s concerns about the Agency’s procurement practices, we took immediate actions. These actions included taking steps to ensure that the Agency’s internal processes meet federal regulations. Additionally, we obtained a review by an outside, independent authority regarding FMCS procurements made over a period of years. We conducted a prompt and thorough investigation and a review of our own internal processes. With the settlement of the employment dispute, the conclusion of our own investigation and reviews by outside authorities, the allegations were dropped and outside authorities indicated they would take no further action."

Subsequently, Congressional committee staff in both the House and Senate looked into the allegations that prompted the articles in the Examiner and concluded their inquiries without making any findings against the agency or its employees.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/


text:
Rajshahi College

Rajshahi College (Bengali: রাজশাহী কলেজ) is a public educational institution in Rajshahi, Bangladesh. Established in 1873, it is the third oldest college in Bangladesh after Dhaka College and Chittagong College. In 1895, Rajshahi College was the first institution in the territories now comprising Bangladesh to award a graduate (master's) degree. The first two master's degree candidates, Cayan Uddin Ahmed (Chawyone Uddin Ahmed) and N. N. Lehari, after graduation became Chief Secretary of Bengal and Session Judge respectively. Later both were awarded Khan Bahadur and Roy Bahadur titles.

Rajshahi College offers three years Degree pass course and four years honours degree courses in various disciplines under the University of Rajshahi. The college is affiliated with the University of Rajshahi. Since 1996, it has stopped enrolling higher secondary students. It again started enrolling higher secondary students in 2010. Situated in the city center, Rajshahi College is adjacent to Rajshahi Collegiate School and is very near the famous Barendra Museum.

History
The origins of the college were in a private English School that had been founded in Rajshahi in 1828 by the concerted efforts of many of the region's most prominent citizens (this school is now known as Rajshahi Collegiate School). In 1836 the school was taken over by the provincial government of Bengal and was converted into a Government Zilla (or District) School. In 1873, again through the concerted efforts and appeals of local citizens, the Zilla School was given the status of an Intermediate College, and F.A. courses were introduced into its curriculum. With further development this college was accorded "first-grade rank" in 1878, which meant that it could teach B.A. courses and be affiliated to the University of Calcutta. The name "Rajshahi College" came with the first-grade rank in 1878. The year 1881 saw the inauguration of the M. A. classes; B.L. classes were added in 1883. The postgraduate Departments in Arts and in Law continued till 1909 when they were withdrawn because the college could not meet the requirements of the New Regulations of the University of Calcutta which came into force in that year.

Starting with only 6 students on the roll in 1873, the college counted 100 in 1878, 200 in 1900, 400 in 1910, 800 in 1920 and no less than 1000 in 1924; there was only one Muslim student at the college in 1873; 5 years later it was still one, but the figure rose to 156 in 1916 and climaxed at 215 in 1924. Subsequently, of course, Muslim numbers rose in the college and after 1947 eventually exceeded the Hindu numbers. Today, in independent Bangladesh, the college has been accorded "University College" status and no longer teaches Higher Secondary or Intermediate courses. Currently, it is part of the University of Rajshahi system of Bangladesh and its curriculum includes only undergraduate and post-graduate courses in a variety of disciplines.

Buildings
The main administrative building (shown in the picture), is a good example of British Indian colonial architecture. Other important older buildings of the colonial period include the Fuller Hostel Biology Building, Chemistry Building, Physics Building, former Muslim Hostel etc. Newer buildings include the Library and Auditorium, an Arts building, both dating from the 1950s and a new Science building, dating from the 1990s.

Rajshahi College and the Language Movement
Immediately after the killing of students in Dhaka on 21 February 1952 students in Rajshahi College built what is often thought to be the first (but short lived) martyr monument dedicated to the Language Movement. The present monument to the Language Movement dates from 1973. It was built to replace an earlier monument, built in 1969, that was destroyed by Pakistani forces in 1971.

E-Service
E-Service plays an important role to bring momentum, transparency and accountability in government service. To achieve the goals of A-2I Programme of Digital Bangladesh, Rajshahi College has taken some initiatives.

There are about 30,000 students pursuing education at HSC, Pass, Honors and Masters levels at Rajshahi College. About 250 teachers are engaged in teaching them and more than 150 other employees support the administration, accounts and all sorts of activities in this college. Frequent change in Principals and weakness in record keeping created serious difficulties in preserving records of essential documents and information of students. Moreover, it is much difficult and a tedious task for the administration to maintain and ensure good governance of the college.

Keeping this in mind, Rajshahi College made an effort to establish exclusively a robust basis for the whole institution and started an ICT program in February 2012 with a view to keeping records of all data, information relating to students, teachers, finance and other things as well. Rajshahi College might be the sole institute which dared to manage its all data and information digitally and has become a pioneer institute to do all its jobs online.

Notable alumni and faculty
Highly distinguished alumni and faculty of Rajshahi College include:
 * Radhabinod Pal
 * Khademul Bashar
 * Enamul Haque, professor from 1948 to 1952
 * Qazi Motahar Hossain
 * Hasan Azizul Huq
 * Akshay Kumar Maitreya
 * Jadunath Sarkar
 * Ashrafuddin Ahmad Chowdhury
 * Maqbular Rahman Sarkar
 * Abdur Rahman Khan, Divisional Inspector School
 * Pramathanath Bishi
 * Pramathanath Bishi

title: Rajshahi College


text:
Minamiechizen, Fukui

Minamiechizen (南越前町) is a town located in Fukui Prefecture, Japan. , the town had an estimated population of 10,745 in 3,470 households and a population density of 31 persons per km². The total area of the town was 343.69 sqkm.

Geography
Minamiechizen is located in central Fukui Prefecture, bordered by Gifu Prefecture to the west, Shiga Prefecture to the south and the Sea of Japan to the west. Parts of the coastal area of the town are within the borders of the Echizen-Kaga Kaigan Quasi-National Park.

Neighbouring municipalities

 * Fukui Prefecture
 * Tsuruga
 * Echizen
 * Ikeda
 * Echizen (town)
 * Shiga Prefecture
 * Nagahama
 * Gifu Prefecture
 * Ibigawa

Climate
Minamiechizen has a Humid climate (Köppen Cfa) characterized by warm, wet summers and cold winters with heavy snowfall. The average annual temperature in Minamiechizen is 13.1 C. The average annual rainfall is 2608.2 mm with December as the wettest month. The temperatures are highest on average in August, at around 25.4 C, and lowest in January, at around 1.7 C. Parts of the town are located within the extremely heavy snowfall area of Japan.

Demographics
Per Japanese census data, the population of Minamiechizen has declined over the past 50 years.

History
Minamiechizen is part of ancient Echizen Province. During the Edo period, the area was mostly part of the holdings of Fukui Domain and Nishio Domain under the Tokugawa shogunate. Following the Meiji restoration, and the establishment of the modern municipalities it was organised into part of Nanjō District in Fukui Prefecture. Minamiechizen was formed on 1 January 2005, by the merger of the two former towns of Imajō and Nanjō, and the former village of Kōno (all from Nanjō District).

Economy
The economy of Minamiechizen is mixed, with agriculture and commercial fishing prominent.

Education
Minamiechizen has four public elementary schools and there middle schools operated by the city government. The town does not have public high school.

Railway
JR West surrendered ownership of the Hokuriku Main Line in Minamiechizen to third-sector company Hapi-line Fukui on 16 March 2024. Hapi-line Fukui
 * Minami-Imajō, Imajō, Yunoo, Nanjō

Local attractions

 * Echizen-Kaga Kaigan Quasi-National Park
 * Somayama Castle, National Historic Site

title: Minamiechizen, Fukui


text:
Newbottle

Newbottle may refer to the following places in England:

 * Newbottle, Northamptonshire
 * Newbottle, Tyne and Wear

title: Newbottle


text:
File:Human Language Families.PNG

Map of Human Language Families

Base: Sprachen der Welt by Stern

title: File:Human Language Families.PNG


text:
Talk:Princeton Engineering Anomalies Research Lab

Exact date of foundation?
Is it possible to specify the exact date of foundation? I can't find anything about it. 2A02:908:375:D00:AC9D:1E94:AB33:D403 (talk) 06:36, 18 April 2023 (UTC)


text: User talk:69.231.152.224

title: User talk:69.231.152.224


text:
Federal Mediation and Conciliation Service

The Federal Mediation and Conciliation Service may refer to either:
 * Federal Mediation and Conciliation Service (Canada)
 * Federal Mediation and Conciliation Service (United States)


text:
Willie Montañez

Guillermo Montañez Naranjo (born April 1, 1948) is a Puerto Rican former professional baseball first baseman, who played in Major League Baseball (MLB) for the California Angels (1966), Philadelphia Phillies (1970–1975, 1982), San Francisco Giants (1975–1976), Atlanta Braves (1976–1977), New York Mets (1978–1979), Texas Rangers (1979), San Diego Padres (1980), Montreal Expos (1980–1981), and Pittsburgh Pirates (1981-1982). He batted and threw left-handed.

Along with Tito Fuentes, Montañez was most noted for being one of MLB's two most flamboyant personalities during the 1970s, so much so that he was labeled a hot dog by detractors. He once commented, “I don't mind being called a hot dog, but it all depends on how people say it."

California Angels
Montañez was born in Cataño, Puerto Rico, and was originally signed by the St. Louis Cardinals as an amateur in 1965. After one season with the Florida Rookie League Cardinals, he was selected by the California Angels in the 1965 Rule 5 draft, and made his major league debut with the Angels in their 1966 season opener as a pinch runner for Norm Siebern in the 14th inning. The Chicago White Sox won the game in the bottom of the inning before Montañez could log an at bat, however, he did have two at bats without a hit with the Angels before being returned to the Cardinals on May 5.

Philadelphia Phillies
On October 7, 1969, the Cardinals traded Curt Flood, Byron Browne, Joe Hoerner and Tim McCarver to the Philadelphia Phillies for Dick Allen, Jerry Johnson and Cookie Rojas. When Curt Flood refused to report to his new team, the Cards sent Montañez and Jim Browning to the Phillies to complete the trade. He appeared in eighteen games with the Phillies in 1970, mostly in the outfield, and collected three runs batted in and scored three runs.

In 1971, Montañez was tabbed as the starting center fielder for the Phillies, and responded with a career-high 30 home runs (still a Phillies' rookie record), 99 runs batted in, and a league-leading thirteen sacrifice flies. He finished second behind Atlanta Braves catcher Earl Williams in the National League Rookie of the Year balloting.

In 1972, Montañez led the National League in doubles (39), and tied for the league lead in outfield assists with fifteen. With Del Unser's arrival in Philadelphia in 1973, Montañez returned to his natural position at first base where he stayed for the remainder of his career.

Montañez batted over .300 for the first time in his career in 1974 (.304). Made expendable after the Phillies claimed Dick Allen off waivers from the Atlanta Braves one week earlier, he was traded to the San Francisco Giants for Garry Maddox on May 4, 1975.

San Francisco Giants
Though he was batting .286 with the Phillies at the time of the trade, he ended the season with a .302 batting average for the second of his three consecutive seasons batting over .300. He also collected a career high 101 runs batted in between his two teams, and emerged as one of the league's top fielding first basemen, leading the league first basemen in assists for the first of three times in his career (1975, 1976 and 1978).

Atlanta Braves
For the second season in a row, Montañez found himself changing teams in the middle of the season when the Giants sent him to the Braves with Jake Brown, Mike Eden and Craig Robinson for Darrell Evans and Marty Perez on June 13, 1976. As a result, Montañez played in an unusual 163 games. At the time of the trade, Montañez had appeared in sixty games for the Giants. He then appeared in 103 games for the Braves, bringing his total to 163 games in a 162-game season. He was named first baseman on The Sporting News NL All-Star team.

The Braves lost 101 games in 1977, and Montañez was selected as the team's sole representative at the 1977 All-Star Game. He went 0 for 2 in his only All-Star game.

New York Mets
Montañez was dealt by the Braves to the New York Mets in the first four-team deal in Major League Baseball history on December 8, 1977 that also involved the Texas Rangers, Pittsburgh Pirates and a total of eleven players changing teams. The Mets also got from the Rangers Tom Grieve and Ken Henderson who was sent to New York to complete the transaction three months later on March 15, 1978. Adrian Devine, Tommy Boggs and Eddie Miller were traded from the Rangers to the Braves. The Rangers received Al Oliver and Nelson Norman from the Pirates and Jon Matlack from the Mets. The Pirates acquired Bert Blyleven from the Rangers and John Milner from the Mets.

Montañez found himself, again, on a last place team as the 1978 Mets narrowly avoided losing 100 games themselves. However, Montañez emerged as one of the few bright spots on the team, leading it with seventeen home runs and 96 runs batted in.

Texas Rangers
Montañez's fortunes with the Mets turned around substantially in 1979. He was batting only .234 with five home runs and 47 runs batted in when the Mets sent him to the Texas Rangers for two players to be named later. In 71 fewer games with the Rangers, he swatted eight home runs and batted .319. Following the season, he was traded to the San Diego Padres for Tucker Ashford, Joe Carroll and future Hall of Famer Gaylord Perry.

Montreal Expos
The 1980 Montreal Expos were in a tight division race with the Philadelphia Phillies throughout the season. They completed a deadline deal for Montañez on August 31, 1980 for Tony Phillips and cash. In fourteen games with the Montreal Expos, Montañez batted .211 with one run batted in.

Return to Philadelphia
On August 20, 1981, he was again involved in a trade with John Milner—this time going from the Montreal Expos to the Pittsburgh Pirates. He was released by the Pirates on July 1, 1982. Ironically, Milner was released by the Expos four days later, and signed with the Pirates later in the month. Montañez signed with the Phillies for the remainder of the season, at the end of which he retired.

Montañez had a career .992 fielding percentage at first base, and led National League first basemen in assists in 1975, 1976 and 1978. From 1975 to 1978 he collected 349 runs batted in.

title: Willie Montañez


text:
User talk:Danny196

Welcome!

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Hi Danny, stuff about you should go on this page, not in the encyclopedia articles. I hope you enjoy contributing to Wikipedia. If you have any questions, feel free to ask me.--Pharos 00:40, 14 Feb 2005 (UTC)

Danny, if you want to put up stuff about your friends up, please put it in your namespace--Pharos 00:54, 14 Feb 2005 (UTC).

Nah Danny, forget Pharos, and keep up the good work. Post some more pictures of you, cause they're kool. --Abdull 22:14, 25 Feb 2005 (UTC)

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title: User talk:Danny196


text:
Talk:List of Nokia products

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title: Talk:List of Nokia products


text:
Night Train Lane

Richard Lane (April 16, 1928 – January 29, 2002), commonly known as Dick "Night Train" Lane, was an American football cornerback who played for 14 years in the National Football League (NFL). He played for the Los Angeles Rams (1952–1953), the Chicago Cardinals (1954–1959), and the Detroit Lions (1960–1965).

As a rookie in 1952, Lane had 14 interceptions, a mark that remains an NFL record over 70 years later. He played in the Pro Bowl seven times and was selected as a first-team All-Pro player seven times between 1956 and 1963. His 68 career interceptions ranked second in NFL history at the time of his retirement and still ranks fourth in NFL history. He was also known as one of the most ferocious tacklers in NFL history and was inducted into the Pro Football Hall of Fame in 1974. He was named to the National Football League 50th Anniversary All-Time Team in 1969, named to the 75th Anniversary All-Time Team in 1994 and unanimously named to the National Football League 100th Anniversary All-Time Team in 2019. In 1999, he was ranked number 20 on The Sporting News' list of the 100 Greatest Football Players.

After retiring from professional football, Lane worked for the Detroit Lions in various administrative positions from 1966 to 1972 and then held assistant coaching positions at Southern University (1972) and Central State University (1973). For 17 years, from 1975 to 1992, he was in charge of Detroit's Police Athletic League.

Youth
Lane was born in Austin, Texas, in April 1928. When he was three months old, he was abandoned by his birth parents, a prostitute and pimp. He was found, covered in newspapers, in a dumpster. Lane later recalled, "My father was called Texas Slim. I never saw him—I don't know if he's the one that told my mother to throw me away. A pimp told my mother I had to go. They put me in a trash can and took off. Some people heard me crying. They thought it was a cat."

Lane was adopted and raised by Ella Lane, who also had four other children. As a youth in Austin, Lane grew up poor, busing tables at local hotels and shining shoes on Congress Avenue. He also helped his mother with a laundry business she ran out of the home. Lane became known as "Cue Ball" and later recalled how he acquired the nickname: "I was in a pool hall in 12th street. We were playing for money, maybe a dime. As soon as I made the eight ball, the other guy took off running. He didn't want to pay. I grabbed that cue ball and just as he made the corner I threw it and hit him upside the head. The guy didn't know what had hit him."

Anderson High School
Lane attended L.C. Anderson High School, Austin's segregated high school for African Americans. He played basketball and football and was a member of the school's 1945 and 1946 football teams. The 1945 team was runnerup in the Prairie View Interscholastic League, an association of black schools in Texas.

Negro league baseball
After graduating from high school, Lane lived for a time in Council Bluffs, Iowa, with his birth mother, Etta Mae King. She had visited during Lane's youth, and the two reconciled. His mother and a man had opened a tavern in Council Bluffs. While in Council Bluffs, a baseball scout signed Lane, and he played for a time with the Negro league Omaha Rockets, a farm team for the Kansas City Monarchs.

Scottsbluff Junior College
In the fall of 1947, Lane enrolled at Scottsbluff Junior College in Scottsbluff, Nebraska. He played one season of college football at Scottsbluff. He was the only African American player on the team, and a clipping from the college newspaper noted, "He is outstanding for his vicious tackles, hard running and pass snatching." The Scottsbluff football team compiled a 5–3–1 record with Lane in the lineup in 1947 and finished in third place in the Nebraska Junior College Conference.

Military service
In 1948, Lane enlisted in the United States Army and served for four years. He served at Fort Ord on Monterey Bay in California and played on a Fort Ord football team. In 1951, he caught 18 touchdown passes for Fort Ord. He received second-team All-Army honors in 1949 and first-team honors in 1951. After his discharge from the Army, Lane worked in an aircraft plant in Los Angeles, lifting heavy sheets of metal out of a bin and placing them into a press.

Signing and nickname
While working at the aircraft plant in Los Angeles, Lane passed the Los Angeles Rams offices on his bus ride to work. He walked into the office with a scrapbook of clippings in 1952 and asked for a tryout. He was recommended to the Rams by Gabby Sims and signed as a free agent. Lane initially tried out as a receiver, the position he had played at Fort Ord, but was switched to a defensive back by the Rams. In the Rams' first scrimmage on August 3, 1952, Lane drew praise as "the outstanding player in the scrimmage by a country mile" due to his "ferocious" approach to the game and his speed in chasing down Elroy Hirsch. After the scrimmage, Rams head coach Joe Stydahar said, "Lane came out here to make the ball club. Well, last night he got himself a job."

Lane acquired the nickname "Night Train" during his first training camp with the Rams. Teammate Tom Fears had a record player in his room and frequently played the record, "Night Train", by Jimmy Forrest. The record was released in March 1952 and was the #1 R&B hit for seven weeks. According to an account published by the Los Angeles Times in August 1952, "Whenever Fears plays it Lane can be found in the hall outside Tom's room dancing to the music." Lane was initially uncomfortable with the racial implication of the nickname, which had been bestowed on him by his white teammates, but he embraced it after a newspaper reported on his performance against Washington Redskins star Choo Choo Justice with the headline, "Night Train Derails Choo Choo".

1952 season
As a rookie in 1952, Lane appeared in all 12 regular season games and broke the NFL single-season record with 14 interceptions. He also led the league with 298 interception return yards and two interceptions returned for touchdowns. In his first NFL game, a 37–7 loss to the Cleveland Browns, Lane was credited by the Los Angeles Times with playing "a positively sensational game at defensive halfback (he made about 50% of the tackles)." On December 7, 1952, he intercepted three passes in a 45–27 victory over the Green Bay Packers, including an 80-yard return of a pass from Tobin Rote. The following week, he intercepted three more passes in a 28–14 victory over the Pittsburgh Steelers, including one that he returned 42 yards for a touchdown. However, he sprained an ankle after making his third interception against the Steelers and was lost to the Rams for their playoff game against the Detroit Lions. The NFL later prepared a list of the greatest single-season performances of all time and ranked Lane's 1952 season fourth on that list.

1953 season
After Lane blocked two field-goal attempts during a July 1953 scrimmage, Rams coach Hamp Pool said, "Night Train has the reflexes of a cat. It just doesn't seem possible that a man can come in from so far out and get in front of the ball in a matter of a couple of seconds." During the 1953 season, Lane appeared in 11 games for the Rams, but he intercepted only three passes. The highlight of his 1953 season was a blocked field goal against the Green Bay Packers; Lane blocked the kick at the Rams' 25-yards line, caught it on the bounce 45 yards downfield, and returned it for a touchdown.

Chicago Cardinals
Lane had signed with the Rams for $4,500. After the two seasons he had with the Rams, he was offered a $2,500 salary increase. He then said it wasn't enough money in his view and that they should trade him. Not long after, in January 1954, the Rams traded Lane to the Chicago Cardinals in a three-team deal that also involved Don Doll. During the 1954 season, Lane appeared in all 12 regular season games for the Cardinals and again led the NFL in both interceptions (10) and interception return yards (181). Lane was occasionally used as a receiver by the Cardinals, and on November 13, 1955, he caught a pass from Ogden Compton, a play that covered 98 yards, the second longest pass in NFL history up to that time.

Lane remained with the Cardinals for six seasons from 1954 through 1959, appearing in 68 games and intercepting 30 passes. During his years with the Cardinals, Lane received All-NFL honors in 1954 (AP and UPI second team), 1955 (UPI second team), 1956 (AP and UPI first team), 1957 (Sporting News first team), 1958 (AP second team), 1959 (NEA first team). He was also invited to play in the Pro Bowl in 1954, 1955, 1956, and 1958.

Detroit Lions
On August 22, 1960, the Cardinals traded Lane to the Detroit Lions in exchange for lineman Gerry Perry. Lions great Joe Schmidt later called it "one of the greatest trades that will ever be made in any sport." At the time of the trade, Lions head coach George Wilson noted: "He has a reputation as a gambler. We are aware of that but he still has speed and experience."

In the Lions' first win of the 1960 season, a 30–17 victory over the Baltimore Colts, Lane intercepted a Johnny Unitas pass and returned it 80 yards for a touchdown, quickly becoming a fan favorite in Detroit. In his first two seasons with the Lions, Lane intercepted 11 passes for 175 return yards. In all, Lane played six seasons with the Lions from 1960 to 1965, appearing in 66 games with 21 interceptions for 272 yards. During his time with the Lions, Lane received All-NFL honors in 1960 (UPI, NEA, and Sporting News first team), 1961 (AP, NEA, and Sporting News first team), 1962 (AP, UPI, Sporting News, and NEA first team), 1963 (UPI and Sporting News first team). He was also invited to play in the Pro Bowl in 1960, 1961, and 1962.

Lane appeared in the 1962 Pro Bowl despite suffering from appendicitis. Weakened and in pain, he blocked an extra point kick and intercepted a Y. A. Tittle pass and returned it 42 yards for the West All-Stars. He checked into a Los Angeles hospital the next day and had his appendix removed.

In early July 1963, Lane married jazz singer Dinah Washington and began serving as her business manager, leading to reports that he might not continue his football career. However, he signed a contract with the Lions in late July. Lane intercepted five passes and recovered two fumbles in 14 games for the 1963 Lions.

Lane was hampered by injuries after the 1963 season. In August 1964, he was injured in a pre-season game, had surgery on his knee, and was out of action for the first part of the 1964 season. Lane ultimately appeared in seven games for the 1964 Lions, managing only one interception, the lowest total of his career up to that point.

On September 7, 1965, after undergoing off-season knee surgery, Lane, at age 37, was released by the Lions. When no other team claimed him, Lane returned to the Lions as a taxi squad player. He was returned to the active lineup on October 20, 1965, appearing in seven games with no interceptions for the first time in his career.

Ferocious tackler
Lane was known as a ferocious tackler, and his style of play led to changes in the rules of the game. In 1961, he tackled Jon Arnett by the face mask as he ran at full speed down the field. Arnett lay motionless on the field after the tackle, and the play left a lasting impression. The following year, the NFL adopted a rule prohibiting the grasping of an opponent's face mask.

Lane's practice of tackling opponents about the head and neck, which was then a legal technique, was sometimes called a "Night Train Necktie". It later became known as a clothesline tackle and prohibited. He later explained the rationale for his practice of necktie tackling:"My object is to stop the guy before he gains another inch. ... [I]f I hit them in the legs they may fall forward for a first down. ... I grab them around the neck so I can go back to the bench and sit down."

In 2009, a film produced by the NFL ranked Lane No. 2 on its list (behind Dick Butkus) of the most feared tacklers in league history. The film also credited Lane's practices with the prohibition of clothesline tackles.

In the book Paper Lion by George Plimpton, former Detroit Lions assistant coach Aldo Forte recalled a hit that Lane placed on then New York Giants quarterback Y. A. Tittle in 1962 that literally "knocked the plays out of his head", rendering the quarterback unable to remember any of the Giants' plays until after halftime.

Career accomplishments and honors
During his 14 years in the NFL, Lane recorded 68 interceptions, 1,207 interception return yards, and five touchdowns on interception. He also recovered 11 fumbles, returning them for 57 yards and one touchdown. From 1954 to 1963, he was selected as a first-team All-NFL player seven times and played in seven Pro Bowls. His single-season record of 14 interceptions still stands despite the lengthening of the NFL season from 12 to 14 to 16 and eventually 17 games. Lane's 68 career interceptions ranked second in NFL history at the time of his retirement and still ranks fourth in NFL history as of the end of the 2023-24 NFL season. His 1,207 interception return yards also ranked second in NFL history when he retired and still ranks sixth in NFL history. His 298 interception return yards in 1952 was three yards short of the NFL record at the time and remains the seventh best single-season total in NFL history.

Lane has received numerous honors for his contributions to the sport. His honors include the following:
 * In September 1969, Lane was one of 16 players named to the all-time All-Pro team selected by the Pro Football Hall of Fame. He was also named to the NFL's 1950s All-Decade Team in August 1969.
 * In February 1974, Lane was named to the Pro Football Hall of Fame. At the induction ceremony in July 1974, Lane was introduced by his high school coach W. E. Pigford. In his speech, Lane spoke out against the NFL's treatment of African Americans as "stepchildren" and added, "I hope the black players will band together to deal with the problem of no black coaches, no black managers and few black quarterbacks in pro football."
 * In January 1988, Lane was named to the Michigan Sports Hall of Fame. At the time, he said, "I was feeling a little put out about having to wait more than 10 years after getting into the pro Hall of Fame to get into the state, but I'm happy that I'm finally in."
 * In August 1994, he was named to the National Football League 75th Anniversary All-Time Team selected by a 15-person panel of NFL and Pro Football Hall of Fame officials, former players, and media representatives. He was the only former Detroit Lions player so honored.
 * In August 1999, Lane was ranked number 20 on The Sporting News' list of the 100 Greatest Football Players, making him the highest-ranked defensive back, the Cardinals' highest-ranked player and the Lions' second highest-ranked player after Barry Sanders.
 * In 2001, Lane was inducted into the Texas Sports Hall of Fame.
 * In August 2006, Lane became one of eight charter inductees into the Arizona Cardinals' Ring of Honor.
 * In 2020, Lane was named to the NFL's 100th Anniversary All-Time Team.

Post-NFL career
After retiring from professional football, Lane worked for the Detroit Lions in various administrative positions. He was the first African American to work in the Lions' front office. In February 1972, Lane quit his job with the Lions to become an assistant football coach at Southern University, a historically black university in Baton Rouge, Louisiana. He left Southern University in June 1973 to become an admissions counselor and assistant football coach at Central State University, a historically black university in Wilberforce, Ohio. In January 1974, he resigned his position at Central State to accept a job in Los Angeles as a bodyguard and personal assistant for television star, Redd Foxx.

In October 1975, Lane was hired to manage Detroit's Police Athletic League. He remained in charge of the program for 17 years and oversaw its expansion to 16 centers with 20,000 participants. He retired from the post at the end of 1992.

Personal life and death
Lane was married three times. He married his first wife, Geraldine Dandridge, in April 1951. The couple separated in August 1962 and was divorced in January 1963.

In July 1963, Lane married jazz singer Dinah Washington at a ceremony in Las Vegas. It was the sixth marriage for Washington and the second for Lane. On December 14, 1963, Lane discovered Washington dead at their home at 4002 Buena Vista Street in Detroit with a bottle of prescription pills on the night stand beside her.

In 1964, Lane married school teacher Mary Cowser, who in 1955 became the first African American woman to appear in Coca-Cola advertisements. They had a son, Richard Ladimir Lane, who was born in 1965. The marriage ended in divorce after ten years.

In 1994, Lane moved from Detroit back to his hometown of Austin, Texas. Due to reduced mobility from diabetes and knee injuries, he spent the last two years of his life at the Five Star Assisted Living facility in North Austin. He died there from a heart attack in January 2002 at age 73, after playing dominoes and while listening to jazz in his room. His family believed that he also suffered from chronic traumatic encephalopathy (CTE) brought on by football-related brain injuries.

title: Night Train Lane