Italian Sponge Cake With Vermouth & Lemon Custard

Total Time
1hr 30mins
Prep 1 hr
Cook 30 mins

A light dessert filled with lemon custard and frosted with sweetened whip cream.

Ingredients Nutrition

  1. Preheat oven to 350F and cut a circle of parchment paper for the bottom of a 10" spring form pan. Prepare the vermouth soak by combining the coffee and vermouth, set aside.
  2. Cake; beat the eggs for 2 minutes on medium speed. Add the sugar and beat on high for 10 minutes, or until the batter has tripled in volume. (The longer the batter is beaten, the better the result). Blend in the oil just until blended. Gently fold in the flour and baking powder just until the batter is free of any lumps, do not over mix.
  3. Pour the batter into the prepared pan and bake for 30 minutes, or until the cake tester comes out clean. Cake will fall slightly once removed from the oven. Leave the cake in the pan until completely cool.
  4. Cut the cooled cake into 2 layers and brush all of the vermouth soak on both of the layers on the cut sides. Using the top of the cake as the bottom layer (the flat bottom for the top), spread the bottom layer with the lemon custard, then top with the second layer. Frost the entire cake with whipped cream.
  5. LEMON CUSTARD; Prepare the whipped topping mix with 1/2 cup of milk and vanilla extract according to the package directions. Prepare the pudding according to the package but using only 1 cup milk. Fold the pudding into the topping mix until completely incorporated.
  6. SWEETENED WHIPPED CREAM; Beat the whipping cream with the 6 tablespoons of sugar until firm peaks are formed when the beaters are lifted, do not overbeat. Refrigerate until ready to use.


BOSTON — A new Massachusetts sales tax on software and computer services, part of a massive transportation financing plan that became law last month, has been met with anger and confusion in the state’s technology sector, prompting calls for its repeal.

Two prominent business-backed organizations, the Massachusetts Taxpayers Foundation and the Massachusetts High Technology Council, are leading an initiative petition that urges lawmakers to repeal the tax when they return from summer recess. At least one bill has already been filed that would do so. If the Legislature doesn’t act, opponents will ask voters to kill the tax on next year’s state ballot.

In an Aug. 14 memo to legislators, the council’s president, Christopher Anderson, said the tax threatens to make the state less competitive at a time when tech companies are expected to lead an economic revival.

The measure imposes the state’s 6.25 percent sales tax on certain computer system design services and the “modification, integration, enhancement, installation, or configuration of standardized or prewritten software.” It’s unclear if the Democratic-controlled Legislature will revisit the issue.

Brewer noted that the tax, originally proposed by Gov. Deval Patrick, is one of several contained in the law that promises $800 million in new revenue for transportation and came in response to concerns from the private sector that the state’s aging infrastructure was stunting economic growth.

For example, customized software that incorporates prewritten software would be taxable, unless the extent of the prewritten material was “inconsequential.” A plug-in that helps create a customized version of a product, such as an Excel spreadsheet, would be taxed. But enhancements made to open source software, generally free and downloadable from the Internet, would be exempt.

The revisions did not appear to be softening opposition to the tax, especially among smaller software services firms that say they’ll be hurt the most.

“It’s a very bad law,” said Andy Singleton, founder and chief executive of Assembla, a 7-year-old Needham company that makes Web-based software. “They can’t narrow it to the point where it makes sense. It’s written to apply to almost anything that a computer programmer does.”

The tax could prompt firms to shed jobs or produce more work outside of Massachusetts, Singleton says. Many of Assembla’s 40 employees already work outside the state or even the U.S., using an online platform to share tasks.


January 11, 2011

Project Maps Out Plan To Save Coral Reefs

The scientists say that reefs are under pressure from a variety of threats, including rising sea temperatures, increased acidity, overfishing and pollution.

The Edge plan focuses on the most evolutionarily distinct and globally endangered species.  The project will take a regional approach to conservation.

The plan would focus on the "coral triangle" around the Philippines, the west Indian ocean around the Mozambique channel, and in the Caribbean channel.

"Coral reefs are threatened with functional extinction in the next 20-50 years, due predominantly to global climate change," Catherine Head, coordinator of the reefs project, said in a press release. "In these regions, we'll be supporting and training in-country conservationists to carry out research and implement targeted conservation actions," she said.

"Their projects will last initially for two years. We provide them with a whole host of tools to carry out their projects including funding and intensive training."

Coral reefs are the planet's most diverse marine ecosystem and they harbor up to a third of all marine life, despite taking up under 0.2 percent of the ocean floor.  Climate change causes corals to bleach.

"Bleaching occurs when sea temperatures rise and this causes the coral tissue to expel their symbiotic algae called zooxanthellae "“ these are what give the coral their color," said Head.

"2010 seems set to have been one of the worst years for coral bleaching. There have been reports on the coast of Indonesia of up to 100% bleaching of many coral colonies. In 1998, 16% of the global coral reefs were killed through bleaching."

A coral cannot photosynthesize when it is bleached.  There is a limited period of time where the coral needs to re-acquire zooxanthellae or else it will die.

"2010 was an El NiÃƒ±o year so sea-surface temperatures were very high "“ but these hot years are coming with increasing frequency now," Rachel Jones, a keeper at London zoo's aquarium, told The Guardian.

Among the 10 species chosen to start the Edge project are the pearl bubble coral, which is a food source for the hawksbill turtle, and the Mushroom coral.

The conservationists said that part of the solution in the future will be to designate more of the ocean as marine protected areas.

The focus of the project will remain on increasing the resilience of reefs to environmental change.

"That means trying to reduce overfishing and pollution pressures," Jones told The Guardian.

"Where we see reefs that are in very good condition, such as the Chagos archipelago where the reef is in fantastic condition despite being hit by catastrophic bleaching in 1998, it recovered better and quicker than anywhere else in the Indian Ocean."

She added: "The environment is changing faster now than it ever has done before. Corals have evolved to live within a very specific set of parameters. They're right at the interface between air and sea and it's a very difficult environment to live in."

"But they've evolved to live there as long as those parameters are steady. At the moment those parameters are shifting in a way that the corals just can't keep up with."

Heather Koldewey, international marine and freshwater conservation program manager for ZSL, said in a statement that "Corals are one of the most threatened groups of animals on our planet and iconic flagships of the marine environment."

On the Net:


Tiny microbes may have been responsible for the largest extinction event our planet has ever seen.

The end-Permian extinction was the most catastrophic mass extinction the Earth has ever seen. It started roughly 252 million years ago — long before the dinosaurs — and it continued for 20,000 years. By the time it was over, nearly 90 percent of all life on Earth had been destroyed, scientists say.

The first clue that microscopic microbes could be involved in the greatest die-off the Earth has ever known came when MIT geophysicist Dan Rothman was looking at how carbon levels grew during this time. "The growth was like what you might see in a real estate bubble, or a financial bubble," he said. "If the C02 came from the sudden combustion of a coal field in Siberia it wouldn't behave this way."

To figure out which microbe might have a hand, Rothman took his research to Fournier, who had published a paper about Methanosarcina in 2008. The paper showed that sometime in the past 400 million years, Methanosarcina was the recipient of a gene transfer that allowed it to produce methane more efficiently than ever before. Fournier worked to date the gene transfer more specifically and found it most likely occurred about 250 million years ago.

Los angeles times


But it's true about Duke from the hoop to the 5

Lyric Meaning

But Duke Da God (a Dipset member) is not lying about his basketball skills, or his car (5-series BMW). He’s an honest fellow


Analysis of the sheep genome

The identification of genes controlling several traits of interest in sheep has been accomplished by positional candidate cloning. In these studies, the trait is first mapped to a specific chromosomal region by linkage analysis, which requires families that are segregating for the trait and for polymorphic markers. Microsatellite markers are usually used for these analyses because of their extensive genetic variability. Once the location of a trait is determined by linkage to the markers, possible candidate genes controlling the trait can be inferred because of their proximity to linked markers. It is not necessary to map all possible genes in sheep for this strategy to be effective. Rather, a subset of genes that are mapped in humans and mice have also been mapped in sheep; these genes serve as “anchors” across the comparative maps of the different species. Further study of these positional candidates has revealed naturally occurring mutations that produce phenotypes that are unique to sheep. Thus the genetic analysis of sheep traits advances knowledge not only in this species but provides critical information for understanding biological pathways in mammalian species.

  • ovine
  • comparative gene mapping
  • positional candidate cloning

The Sheep Genome Map

The diploid number in domestic sheep (Ovis aries) is 54, with 26 pairs of autosomes and two sex chromosomes. Ovine autosomes are telocentric with the exception of three pairs of large metacentric chromosomes (1, 2, and 3). The X is the largest acrocentric chromosome, and the Y is a very small metacentric chromosome, which usually looks like a small, square dot. The X chromosome can be recognized in unbanded preparations because of the presence of small short arms, but the telocentric autosomes cannot be distinguished without differential staining techniques (7). In 1995, a standard G-band karyotype for the sheep was agreed upon at the 9th North American Colloquium on Domestic Animal Cytogenetics and Gene Mapping (7). The ovine and bovine standards were correlated based on similar band patterns and mapped gene loci. G-, Q-, and R-banded karyotypes and idiograms for sheep chromosomes have also been presented (1). This report included chromosome-specific molecular markers that had been mapped by in situ hybridization to 24 of the 26 ovine autosomes.

The three metacentric chromosomes in sheep, OOV1, OOV2, and OOV3, are equivalent to centromerically fused bovine chromosomes 1 and 3 (BTA1 and BTA3), BTA2 and BTA9, and BTA5 and BTA11, respectively. These fusions at the centromere are supported by marker order on the sheep and bovine linkage maps (22, 24). Sheep chromosome 9 (OOV9) appears to be the product of a translocation of the telomeric end of BTA9 to BTA14 (22). The remainder of BTA9 corresponds to OOV8.

Development of the linkage map for sheep has been critical for genomic studies in this species. Prior to 1994, only 17 markers were assigned to 7 syntenic groups (7). In 1994, 19 linkage groups containing 52 markers including microsatellites and candidate gene restriction fragment length polymorphisms (RFLPs) were identified (21). These assignments were a consequence of a genome scan initiated to map the Booroola fecundity gene (64) using 12 pedigrees segregating for the Booroola gene. A more extensive ovine linkage map was published by this group in 1995 (22) and contained 246 markers (86 ovine microsatellites, 126 bovine microsatellites, 1 deer microsatellite, and 33 known genes), with marker spacing between 10 and 30 cM across the chromosomes. Total coverage of the map was 2,070 cM (∼75% of the genome), and markers were assigned to all 26 sheep autosomes. This map was constructed using the AgResearch International Mapping Flock (IMF). Three years later, a second-generation ovine genetic map was published by the United States Department of Agriculture (USDA) Agricultural Research Service (ARS) group in Nebraska (24). The map contained 519 markers (402 bovine microsatellites, 101 ovine microsatellites, and 16 known genes) and spanned 3,063 cM across the autosomes, with an average marker spacing of 6.5 cM ( A third generation map has been recently developed (58). This map contains 1,062 loci (941 anonymous loci and 121 gene) and is a compilation of genotype data generated by 15 laboratories using the IMF population (∼jillm/jill.htm). The map spans 3,400 cM (sex-averaged) for the autosomes and 132 cM (female) on the X chromosome.

It should be noted that these ovine linkage maps contain relatively few expressed genes (∼120) because of the difficulty in identifying allelic variation needed for linkage analyses. Additional assignments for ∼250 genes have been made using somatic cell hybrids (8, 82) and in situ hybridization.

There are some estimates of gene order for sheep and cattle, two ruminant species within Bovidae. In one comparison (24), marker order was inverted eight times (>2-cM intervals) when the sheep map was compared with the cattle linkage map (48). Length of the inversions ranged from 3 to 57 cM. This group also found six microsatellites that mapped to sheep linkage groups other than what would be expected based on the bovine homolog. The authors suggest that these may be the result of amplification of a nonhomologous sequence by heterologous primers.

Conserved chromosomal segments between sheep and humans have been determined using human chromosome painting probes on R-banded sheep chromosomes (44) and through the use of human and sheep chromosome painting probes on the Indian muntjac deer (9). A total of 48 human segments were found in sheep chromosomes.

An informational database that includes mapped loci in sheep is now available through the Roslin Institute (United Kingdom) at or Texas A&M University (USA) at The current version of SheepBase contains about 1,492 loci gathered from 507 publications. There are about 369 designated genes, 1,098 type II loci, and 25 other loci (such as blood group polymorphisms) in the database. Of the 2,257 map assignments, 1,413 were based on linkage studies, and 844 were assigned using cytogenetic methods.

Identification of QTLs for Wool, Meat, and Milk in Sheep

Genome scanning projects have been initiated for identification of genes controlling many quantitative traits in sheep. Preliminary reports of the identification of sheep quantitative trait loci (QTLs) include parasite resistance (4, 20), wool production traits (47, 73), milk traits (27), and fecal soiling of the wool or dagginess (56). Development of additional resource flocks is ongoing for traits in sheep such as parasite resistance, out-of-season breeding, carcass traits, and wool production.

Parasite resistance.

Genetic variation between breeds of sheep in levels of resistance to internal parasites has been well documented (5). Thus the development of genetic markers for parasite resistance has received major attention in the last 5 years. Resource families suitable for the identification of genetic markers for resistance to Trichostrongylus colubriformis have been established by Commonwealth Scientific and Industrial Research Organisation (CSIRO, Australia), AgResearch (New Zealand), and the University of Melbourne and for Haemonchus contortus by Louisiana State University, CSIRO, and International Livestock Research Institute (ILRI, Kenya). The initiation of whole genome scans has been reported by CSIRO (4) and AgResearch (20), but information on specific chromosomal regions that contain QTLs for parasite resistance are not yet available.

Wool fiber in sheep is a highly organized structure, with about 90% of the fiber made up of keratin intermediate filament (IF) and keratin-associated proteins (KAP). The keratin IF proteins form 8- to 10-nm diameter filaments that are embedded in a matrix of KAPs. The two types of keratin IF, type I and type II, are paired to form the basic unit of the filament. Wool keratin IF type I genes are 4–5 kb with 6 introns, whereas the type II genes are 7–9 kb with 8 introns (76). In sheep, the type I (KRT1.n) and type II (KRT2.n) genes map to 11q25-q29 and 3q14-q22, respectively (39). These physical map assignments have been confirmed through linkage analysis (63), with KRT1.2 mapping to OOV11 and KRT2.10 and KRT2.13 closely linked on OOV3. A 100-kb DNA fragment containing the keratin type II region of sheep has been characterized (77, 78). Three wool and three wool-related type II genes are found within the region in a tight cluster, which is a common feature of the keratin genes.

The KAPs have been grouped (76) into high-glycine-tyrosine KAPs (encoded for by the KAP6.n multigene family, KAP7, and KAP8), the high-sulfur KAPs (encoded for by the KAP1.n, KAP2.n, and KAP3.n multigene families), and the ultra-high-sulfur KAPs (encoded for by the KAP4.n and KAP5.n multigene families). The KAP genes are small, between 0.6 and 1.5 kb in size, and do not contain introns. The AgResearch IMF families have been used in a linkage analysis to map KAP6.1, KAP7, and KAP8 to OOV1; KAP1.1, KAP1.2, KAP1.3, and KAP3.2 to OOV11; and KAP5.1 to OOV21 (63). KRT1.2 was also linked to the KAP genes on OOV11. Another wool follicle gene, trichohyalin (THH), was assigned to OOV1 as well.

Studies of the expression and regulation of hair and wool keratin genes are ongoing. Several consensus sequences have been identified within the promoters of these genes, including binding sites for AP1 and AP2, two common transcription factor complexes, and for lymphoid enhancer factor 1 (LEF1) (76).

Many phenotypic traits associated with the color and quality of wool have been reported, such as albino, lustrous wool, silky wool, and fibre loss (reviewed in Ref. 89). The genetics of these phenotypes are well characterized, but the actual genes that are responsible are not yet known. It is likely that there is involvement of the keratin IF and KAP genes. Interestingly, two wool mutations, HH1 and HH2, affect not only the amount of medullated hair found within the fleece but also have a pleiotropic effect on the formation of horns. While horns and wool are visually different, both contain substantial amounts of keratin.

Limited reports of genes associated with wool production traits are available. Linkage between KAP6 and wool fiber diameters has been found (73). In addition, a resource population for wool production QTLs has been established (47) using a Merino × Romney backcross. While successful identification of QTLS for yield, standard deviation of follicle diameter in a staple, bulk, clean fleece weight, and greasy fleece weight was reported by these authors, no specific chromosomal regions were indicated.

Fecal soiling.

Dagginess is a trait unique to sheep in which there is excessive fecal soiling of the breech area. This soiling occurs because of scouring and too much wool in the anal region. There is an economic incentive to reduce the incidence or severity of dags, because soiled wool must be discarded or discounted and because fecal matter may contaminate the carcass at slaughter. Preliminary evidence for a QTL for dagginess has been reported (56).

Several QTL projects in sheep are now underway. However, production of suitable resource families for QTL searches has required a minimum of 3–5 years. Thus only preliminary results are available for many of these projects. Full reports of map assignments for these QTLs are forthcoming, but because of the paucity of sheep flocks suitable for QTL studies, only a few of these putative QTLs will be replicated in other studies. The identification of genes associated with ovine QTL will likely be dependent on the study of candidate genes revealed in other species.

Identification of Single Gene Traits in Sheep

Chromosomal assignment of many single-gene traits in sheep are known, including the Booroola fecundity gene (OOV6; 64, 65), the Inverdale fecundity gene (X; 23), callipyge muscle hypertrophy (OOV18; 15), Carwell muscle hypertrophy (OOV18; 61, 69), Belgium Texel double muscling (OOV2; 59), spider lamb syndrome (OOV6; 18), and horns (OOV10; 66).

Booroola fecundity.

The Booroola fecundity mutation, FecBB, is codominant for ovulation rate and partially dominant for litter size, with one copy of FecBB increasing litter size by approximately one to two extra lambs (75). In 1993, FecB was linked to markers from a region of human chromosome 4q (64). Subsequent mapping experiments assigned the locus to a position on ovine chromosome 6 (65), corresponding to human chromosome 4q21–25 (67, 96). Upon investigation of positional candidates from 4q21–25, these two groups independently identified a nonconservative mutation (Q249R) in the intracellular signaling domain of bone morphogenetic protein IB receptor (BMPR-IB), a member of the transforming growth factor-β (TGF-β) receptor family (67, 96). This substitution was in total linkage disequilibrium with FecB.

Growth differentiation factor-5 (GDF5) and BMP4, two natural ligands of BMPR-IB, normally inhibit the secretion of progesterone by sheep granulosa cells, which is necessary for differentiation of granulosa cells and maturation of ovulatory follicles. Granulosa cells from FecBB/FecBB females secreted significantly more progesterone than cells of FecB+/FecB+ females when cultured in vitro (67). These findings suggest that FecBB/FecBB cells are less responsive to the inhibitory effects of GDF5 and BMP4 because of altered functionality of the BMPR-IB receptor, leading to advanced differentiation of the granulose cells and enhanced follicular development in animals carrying the mutation.

Inverdale fecundity.

The Inverdale fecundity gene, FecX, was identified in a prolific New Zealand sheep family descended from a Romney female that produced 33 lambs in 11 lambings (23). Progeny testing suggested that a major gene for prolificacy was located on the X chromosome. Females that are carriers for the Inverdale allele, FecXI, have an increase in ovulation by about one extra egg and give birth to ∼0.6 extra lambs. Females that are homozygous for the gene are infertile and have small nonfunctional “streak” ovaries that are about one-eighth the size of normal ovaries. Males with the Inverdale allele have normal fertility.

To more precisely map the location of the Inverdale locus, 14 microsatellite markers and 7 genes were incorporated into a linkage map of the ovine X chromosome (34), with a recombination length of 141.9 cM. The Inverdale gene was then localized to a 10-cM region on this chromosome (33), corresponding to human chromosome Xp11.2–11.4. A member of the TGF-β superfamily, bone morphogenetic protein 15 (BMP15), also known as GDF9B, is located in this region. The BMP15 protein is specifically expressed in oocytes, but its function is not yet known. Mutations in this gene are in linkage disequilibrium with FecXI, as well as another X-linked hyperprolific trait, Hanna or FecXH (35). The FecXI mutation is associated with a V31D substitution, resulting in a nonconservative change in a highly conserved region of the protein. A single C-to-T transition at nucleotide position 67 of the mature peptide introduces a premature stop codon in the FecXH allele. Both changes would be expected to produce a biologically inactive form of BMP15. It should be noted that these two X-linked fecundity traits as well as Booroola prolificacy are due to mutations in genes from the TGF-β pathway.

The callipyge gene is a mutation in sheep responsible for pronounced muscle hypertrophy of the fast twitch muscle fibers (Fig. 1), primarily in muscles of the pelvic limb (Fig. 2; 10, 50). Interestingly, the hypertrophy is absent at birth and develops only after ∼3 wk of age. Callipyge animals produce leaner, higher yielding carcasses (46, 50), but there is some concern with decreased tenderness of the loin (49, 50). Several postmortem tenderization methods are effective in improving the tenderness of callipyge lamb meat (17).

Fig. 1.

Micrographs of longissimus muscle of normal (A) and callipyge (B) lambs stained for ATPase and NADH tetrazolium reductase activity. Cellular hypertrophy is evident in the callipyge muscle, especially in the fast-twitch, glycolytic fibers, which appear clear to lightly stippled.

Fig. 2.

Conformation of a normal-appearing homozygous callipyge ram (far left) and four callipyge offspring.

Genetic characterization of the locus has demonstrated a unique mode of inheritance termed “polar overdominance” (16, 32), in which only heterozygous offspring inheriting the mutation from their sire express the phenotype. The three other genotypes are normal in appearance. Progeny data indicate that reactivation of the maternal callipyge allele occurs after passage through the male germ line, although this reactivation is not absolute (16).

Our earlier work (15) localized callipyge to the distal end of ovine chromosome 18, based on linkage to markers TGLA122, CSSM18, and GMBT16. The position of callipyge was then more precisely mapped to a 4.6-cM interval between IDVGA30 and OY3 by breakpoint mapping using lambs produced in crosses of the four callipyge genotypes and assuming a polar overdominance model (87). Isolation of eight new microsatellites from bovine (87) and ovine (86) contigs allowed us to more precisely localize callipyge within a 450-kb chromosome segment (6).

To characterize this region, two ovine bacterial artificial chromosome (BAC) clones and a long-range PCR product connecting these two BACs were completely sequenced. The resulting sequence traces of 248,602 bp were aligned with the corresponding human genomic sequence obtained from GenBank (accession numbers AL132711, AL117190, and AL132709), with an overall similarity of 44.7%. Six genes were predicted to reside within the analyzed segment. Two of these genes, DLK1 and GTL2, have been previously reported in humans (83, 97), mice (84, 8, 88), and cattle (30, 31), whereas the remaining genes, DAT, PEG11, antiPEG11, and MEG8, are previously undescribed (12).

Using primers designed from each of these genes, Charlier et al. (12) performed RT-PCR on several tissues sampled from an 8-wk-old lamb, demonstrating that all six transcripts are preferentially expressed in skeletal muscle. Because the corresponding regions in humans and mice contain genetically imprinted loci, the imprinting status of these six genes was investigated using single nucleotide polymorphisms (SNPs). Genomic DNA of lambs contained both alleles of the SNP; however, because all six genes were imprinted, only one allele of each gene was detected in skeletal muscle mRNA. By tracing the alleles back to the lambs’ parents, maternal or paternal imprinting status of each gene was determined. These results demonstrated that DLK1, DAT, and PEG11 are paternally expressed, whereas GTL2, antiPEG11, and MEG8 are maternally expressed (12).

In a subsequent study (13), poly(A)+ RNA was isolated from the longissimus dorsi of 8-wk-old animals of the four callipyge genotypes. Northern blotting and/or RT-PCR were used to examine expression of DLK1, GTL2, PEG11, and MEG8 with G3PDH expression serving as a control for similar amounts of mRNA. These experiments demonstrated that the callipyge mutation does not alter the imprinting status of the four genes but dramatically enhances their expression level in cis (Fig. 3). The paternally expressed genes (DLK1 and PEG11) had increased expression in the C/C and CP/NM genotypes (where P and M superscripts refer to the paternal and maternal chromosomes, respectively), those genotypes that have the paternally expressed callipyge chromosome. For the maternally expressed genes (GTL2 and MEG8), increased expression occurred in the C/C and NP/CM genotypes, which have the callipyge mutation on their maternal chromosome. Thus the callipyge mutation appears to modify the activity of a common regulatory element, which could be either an enhancer (boosted by the callipyge mutation) or a silencer (inhibited by the callipyge mutation). This altered regulatory element may be responsible for the enhanced expression of DLK1, GTL2, PEG11, and MEG8, and possibly other untested genes as well. It is likely that increased expression of one or more of these genes initiates the manifestation of the callipyge phenotype. However, the actual callipyge mutation is not yet known.

Fig. 3.

A model for the callipyge locus.

Rib-eye muscle.

Characterization of another gene responsible for an increase in the rib-eye muscle of lambs has been reported (2). Effects of the rib-eye muscle (REM) locus are less dramatic than for callipyge, with an 11% increase in muscle mass limited to the longissimus dorsi. The REM locus appears to act as a dominant gene with no evidence of imprinting and has only minor effects on meat tenderness (62). Interestingly, the REM locus has been localized to the distal end of ovine chromosome 18 (61, 69). However, recent information (McLaren R, personal communication) indicates that the REM locus lies centromeric to callipyge, suggesting that these traits are not allelic to each other but rather may be controlled by two closely linked genes.

Muscle hypertrophy.

Texel sheep are characterized by generalized muscular hypertrophy (11). Based on the association between mutations in the myostatin gene and double muscling in cattle, a study was recently initiated to investigate the possible involvement of myostatin in the hypertrophy of Belgian Texel sheep (59, 60). In this study, the entire coding sequence of myostatin was determined for the Texel and normally muscled Romanov controls; however, no sequence differences were identified. Interestingly, analysis of microsatellite markers flanking myostatin on chromosome 2 in a (Texel × Romanov) × Romanov backcross pedigree revealed significant linkage (Marcq F, personal communication). Thus the causative mutation responsible for hypertrophy found in Texel sheep may reside outside the coding segments of the myostatin gene.

Spider lamb syndrome.

Ovine hereditary chondrodysplasia or “spider lamb syndrome” (SLS) is a recessive genetic disorder (90) causing skeletal deformities in newborn lambs. Structural abnormalities include abnormally long, spider-like legs, humped and twisted spines, deformed ribs and sternebra, facial deformities, lack of body fat and underdevelopment of muscle (Fig. 4). Multiple irregular islands of ossification are found in shoulders, elbows, and sternum of spider animals upon radiological evaluation (81, 93). It is believed this disorder arose as a mutation in a Suffolk genetic line that was used heavily during the late 1960s because of desirable production and show-ring characteristics. The syndrome has since surfaced in several sheep breeds within the last two decades, including North American Suffolks and Hampshires, and US Southdowns, Shropshires, and Oxfords. In addition, there are reported cases of SLS in New Zealand and Australia, after the importation of several US Suffolks into Australia in 1987 (74).

Fig. 4.

Conformation of a spider lamb (left) compared with a normal flock mate (right), both 1 mo of age. Note the abnormally long legs of the spider lamb.

A project to identify a genetic marker for SLS was initiated at Utah State University in 1991. Genetic mapping of the SLS locus was performed by linkage analysis between microsatellite loci in two large pedigrees produced from matings of two carrier rams to 65 carrier and normal females, who produced 223 normal and 80 spider lambs. After screening 174 markers, results from microsatellite marker OarJMP8, located on the distal end of ovine chromosome 6 (55), revealed significant linkage with the SLS trait (18). A combined LOD (logarithm of the likelihood ratio) of 2.85 at 28.1% recombination was obtained across both pedigrees. Three additional markers that had been previously mapped to this region were also linked to SLS. Final order of the loci in this region was determined as OarJMP8-McM214-OarJMP12-BL1038-SLS. A maximum LOD score of 12.89 was obtained between the SLS locus and BL1038 at 3% recombination.

A comparative mapping approach was initiated to identify candidate genes that are located on ovine chromosome 6 (OOV6). Prior mapping of structural genes on OOV6 indicated that it is orthologous to human chromosome 4 (HSA4; 53) and that genes mapping within HSA4p16.3 are distal to microsatellite markers OarJMP8 and OarJMP12 (Lord et al., Ref. 53). Contained within HSA4p16.3 is the gene for fibroblast growth factor receptor 3 (FGFR3; 91). Mutations in the FGFR3 gene of humans result in varying forms of dwarfism (94). Two groups have produced mice in which the FGFR3 gene is disrupted by homologous recombination (19, 25). Remarkably, mice homozygous for the FGFR3 knockout display skeletal abnormalities that are similar to SLS. From these knockout studies, the function of FGFR3 can be defined as a negative regulator of bone growth. To investigate the possible role of FGFR3 in SLS, a single-stranded confirmational polymorphism (SSCP) analysis of the gene was undertaken. An SSCP spanning from the 3′ end of exon 14 through the 5′ end of exon 18 was predictive of SLS genotype; linkage analysis of this polymorphism in a subset of the pedigrees resulted in a LOD score of 18.58 at 0% recombination. To further investigate FGFR3, genomic DNA from one homozygous normal animal, two carriers, and two spider lambs was amplified and sequenced. Analysis of the sequence revealed a T-to-A transversion in exon 17, resulting in a nonconservative amino acid substitution of valine to glutamine at position 700 within the second tyrosine kinase domain (3). Using a PCR-RFLP designed to detect this polymorphism, researchers have performed population studies including more than 2,000 sheep of differing SLS genotypes suggesting that this is the causative mutation in SLS. It is most likely that the mutation leads to loss of receptor function in homozygotes, resulting in poorly controlled chondrocyte differentiation. There is also a suggestion that heterozygous animals have increased bone growth, making them taller in stature than their homozygous normal counterparts. This might be an explanation for the high frequency of the SLS allele in show-ring animals (Beever JE, unpublished data). Further studies of in vitro receptor function are being conducted.

There are at least three genetic systems controlling development of horns in sheep. In some sheep strains, the halo hair (HH1) locus controls the sex-limited expression of horns and has marked pleiotropic effects on fleece characteristics (28), with the expression of horns associated with highly medullated hairlike fleece. Sheep carrying the HH1N allele produce hair in addition to wool on their backs as young lambs. A second genetic system for horns is expressed in Merino sheep and controlled by the Ho locus, with three alleles: HoP, Ho+, and Hoh1 (52). Polledness (absence of horns) is produced by the allele HoP, which is incompletely dominant in rams and almost completely dominant in females. Ho+ produces horns in both sexes, and Hoh1 produces sex-limited horns. There can also be modification of the growth of horns in intact males by the autosomal scurs locus (52).

In an experiment to map the ovine Ho locus (66), pedigrees were established by crossing horned Merino rams with polled New Zealand Romney females. The resulting F1 rams were then backcrossed to Merino females, and horn data were collected on 183 offspring. It was expected that markers on ovine chromosome 1 (OOV1) would be linked to the Ho locus because of the previous assignment of the Bos taurus polled locus to bovine chromosome 1 (BTA1; 36), which is the evolutionary homolog to OOV1. Although 14 markers from OOV1 were tested, no linkage was detected, and OOV1 was excluded. Instead, significant linkage was found with markers on OOV10, and the Ho locus was mapped to the telomere of this chromosome (66). The authors suggest that the Ho locus may correspond to the african horn locus in cattle (36), which has not been mapped, whereas the ovine HH1 locus may be orthologous to the cattle polled locus on BTA1.

The genetic inheritance of many other traits in sheep has been characterized (reviewed in Ref. 52). However, the chromosomal assignment and/or causative mutation is not yet known.

Genes for Economically Important Traits In Sheep Are Isolated Through Functional Cloning

Coat color.

The control of coat color in mammals is based on the ability of melanocyte cells to produce melanin pigments and move the melanin into hair, skin, and wool. Melanin can be either eumelanin (black or chocolate brown, mostly composed of tyrosine) or phaeomelanin (reddish-brown or yellowish tan, mostly composed of cysteine). Melanocytes can produce both eumelanin and phaeomelanin but usually only one pigment type at a time. Binding of α-melanocyte-stimulating hormone (α-MSH) to MSH receptor (MC1-R) on melanocyte cell surfaces initiates production of eumelanin. Absence of the α-MSH signal results in phaeomelanin production. Any number of possible alterations to the normal functioning of this complex system will result in hair, skin, or wool without pigmentation or a dilution of pigmentation. For instance, white color can be caused by several reasons such as the lack of melanocytes or decreased effectiveness of melanin production (89). Another example is the dominant dark coat color in which a mutation in MC1-R causes activation of this receptor even in the absence of α-MSH (54). Recently, the association of a mutation in MC1-R and the dominant black coat color in sheep has been reported (92). In this study, the complete coding region of the MC1-R gene was sequenced from one white and one black sheep of the Norwegian Dala breed. Sequence analysis revealed two mutations in the black animal, resulting in amino acid substitutions M73K and D121N. Design of a PCR-RFLP allowed detection of the mutations in a Norwegian Dala pedigree. All animals (9 black and 13 white) were descendants of a single black female. Both mutations showed complete linkage with black coat color. Black animals were heterozygous for the mutations, whereas the white sheep were homozygous wild type, thereby supporting the dominant inheritance of the black coat color. To further characterize these mutations, they were introduced into the mouse MC1-R gene, and activity of the expressed receptor was assessed using pharmacological assays (92). The M73K mutation was able to constitutively activate the receptor, whereas the D121N mutation did not. When the mutations were introduced together, activation of the receptor still occurred but was less intense. Taken in total, these experiments suggest that the black coat color in sheep is due to the M73K mutation in the MC1-R gene. The mechanism by which M73K activates the receptor is distinct from that in mice and the Alaska silver fox.

Transmissible spongiform encephalopathies (TSE) are characterized as transmissible neurodegenerative disease in mammals, with susceptibility to infection modulated by host-genetic interactions (72). A unique characteristic of TSE, whether sporadic, dominantly inherited, or acquired by infection, is the accumulation of an abnormal, protease-resistant prion protein (PrPSc) in the brain of TSE-infected animals (80). The normal cellular form of the PrP protein (PrPC) is encoded by a host autosomal gene; it has been suggested that the abnormal form (PrPSc) serves as a template for posttranslational misfolding of the normal protein (79).

The TSE of sheep is called scrapie (40). Typical clinical signs of scrapie in sheep begin with mildly impaired social behavior, followed by locomotor uncoordination or ataxia with trembling (26). Pruritis can result from the animal attempting to relieve what seems to be an intense itching by scratching against fence posts or by biting the affected area, and these clinical signs can last from 2 wk to 6 mo. Brain lesions develop late in the incubation period and include neuronal degeneration, vacuole formation, and astroglial cell proliferation.

It has been proposed (41, 43) that the genetic control of susceptibility to scrapie is either closely linked to, or identical to, the prion protein (PRPN) gene. The ovine PRPN gene contains three exons and is over 20 kb (95). The PRPN open-reading frame is contained in exon 3. Polymorphic variants in the coding region of the ovine PRPN gene have been associated with the incidence of both experimental and naturally occurring scrapie (14, 43). In many breeds, including the Cheviot, Swaledale, Ile de France, Shetland, Scottish Halfbred, and Bleu du Maine, the PRPN allele encoding valine at codon 136 (V136) is associated with an extremely high risk of scrapie, with most scrapie-positive sheep of these breeds, either naturally or experimentally infected, possessing the V136 codon (14, 38, 41, 42, 51, 57). However, the V136 variant is rare in British and US Suffolk (38, 41, 70, 95) and at a very low frequency in Japanese Suffolk (45). In contrast to other breeds, variation at codon 171 has been associated with scrapie susceptibility in Suffolk (95). Three amino acid variants at codon 171 [glutamine (Q171), arginine (R171), and histidine (H171)] have been identified (38, 95). In previous studies (70, 95), Suffolk sheep that were scrapie positive (n = 61) were all homozygous for glutamine at codon 171 (QQ171). Also, sheep homozygous for arginine (RR171) or heterozygous (QR171) did not develop scrapie from either natural or experimental challenge in several studies (38, 51, 70, 71, 95). These analyses suggest a relationship between the PRPN QQ171 genotype and scrapie susceptibility in Suffolk sheep. However, this association is not absolute, as 2 of 64 scrapie-positive animals were QR171 in one study (43) and a scrapie-positive Suffolk sheep with the RR171 genotype was found in Japan (45).

An extensive study of a scrapie epidemic in a closed flock of Romanov has been presented (29). Over a 4-year period, 1,015 animals were exposed to scrapie, resulting in 304 deaths. The most resistant PRPN allele was ARR (codons 136/154/171), with no ARR carriers developing scrapie during the epidemic. The VRQ allele was highly susceptible, with 76% of the scrapie-positive animals possessing at least one copy of this allele. Other alleles were ranked by the authors for scrapie resistance/susceptibility. The authors suggest that selection for ARR or AHQ carriers may be a method for increasing scrapie resistance within a flock. However, they note that these animals may still be susceptible to other scrapie strains, and it is possible that these healthy animals may transmit the scrapie agent.

For some ovine traits such as coat color, comparison to similar traits in other species has led directly to the appropriate gene in sheep. This candidate approach at the protein level has also allowed characterization of many other genetic traits in sheep, particularly disease disorders (68). In other cases, such as SLS, localization of the locus through a genome scan narrowed the search for a functional candidate to a specific chromosomal region. However, for traits unique to the sheep, such as Booroola fecundity, callipyge, REM, dagginess, and horns, the candidate gene approach has not been effective because the gene responsible for a similar trait in other species has not yet been identified or characterized. Researchers are seeking to identify these genes in sheep using positional cloning techniques, thereby leading to the discovery of genetic mechanisms not yet explored. Clearly, the analysis of traits unique to sheep has extended our understanding of the functional role and regulation of genes beyond what was known in mice or humans.

  • Article published online before print. See web site for date of publication (

    Address for reprint requests and other correspondence: N. E. Cockett, Utah State Univ., 4700 Old Main Hill, Logan, UT 84322-4700 (E-mail:

View Abstract


File:ScythianC14AsiaEuropeFig6SketchEn 3dGraph.gif|thumb +Search for Videos

Other uses
digitus I manus
digitus primus manus

Bones of the thumb, visible at left.
Princeps pollicis artery+
Dorsal venous network of hand+
Dorsal digital nerves of radial nerve+, proper palmar digital nerves of median nerve+
Infraclavicular lymph nodes+

The '''thumb''' is the first digit+ of the hand+. When a person is standing in the medical anatomical position+ (where the palm is facing to the front), the thumb is the outermost digit. The Medical Latin English noun for thumb is pollex+ (compare ''hallux+'' for big toe), and the corresponding adjective for thumb is pollical+.

# Any of the five digits.

Linguistically, it appears that the original sense was the broader of these two: ''penkwe-ros'' (also rendered as ''penqrós'') was, in the inferred Proto-Indo-European language+, a suffixed form of ''penkwe'' (or ''penqe''), which has given rise to many Indo-European+-family words (tens of them defined in English dictionaries) that involve or flow from concepts of fiveness.

* Having a skeleton of phalanges+, joined by hinge-like joints that provide flexion toward the palm of the hand
* Having a "back" surface that features hair and a nail, and a hairless palm-of-the-hand side with fingerprint+ ridges instead

* Is opposable to the other four fingers
* Has two phalanges rather than three
* Has greater breadth in the distal+ phalanx than in the proximal+ phalanx
* Is attached to such a mobile metacarpus+ (which produces most of the opposability)

Primatologists and hand research pioneers John+ and Prudence Napier+ defined opposition as: "A movement by which the pulp surface of the thumb is placed squarely in contact with - or diametrically opposite to - the terminal pads of one or all of the remaining digits." For this ''true'', pulp-to-pulp opposition to be possible, the thumb must rotate about its long axis (at the carpometacarpal joint+). Arguably, this definition was chosen to underline what is unique to the human thumb.

Anatomists and other researchers focused exclusively on human anatomy, on the other hand, tend to elaborate this definition in various ways and, consequently, there are hundreds of definitions. Some anatomists restrict ''opposition'' to when the thumb is approximated to the fifth digit (little finger) and refer to other approximations between the thumb and other digits as ''apposition''.
Other researchers use another definition, referring to opposition-apposition as the transition between flexion-abduction and extension-adduction; the ''side'' of the distal thumb phalanx thus approximated to the palm or the hand's radial side (side of index finger) during ''apposition'' and the ''pulp'' or "palmar" side of the distal thumb phalanx approximated to either the palm or other digits during ''opposition''.

Moving a limb back to its neutral position is called reposition and a rotary movement is referred to as circumduction+.

The skeleton of the thumb consists of the first metacarpal bone+ which articulates proximal+ly with the carpus+ at the carpometacarpal joint+ and distal+ly with the proximal phalanx+ at the metacarpophalangeal joint+. This latter bone articulates with the distal phalanx+ at the interphalangeal joint+. Additionally, there are two sesamoid bone+s at the metacarpophalangeal joint.

The muscles of the thumb can be compared to guy-wire+s supporting a flagpole; tension from these muscular guy-wires must be provided in all directions to maintain stability in the articulated column formed by the bones of the thumb. Because this stability is actively maintained by muscles rather than by articular constraints, most muscles attached to the thumb tend to be active during most thumb motions.

A ventral forearm muscle, the flexor pollicis longus+ (FPL) originates on the anterior side of the radius+ distal to the radial tuberosity+ and from the interosseous membrane+. It passes through the carpal tunnel+ in a separate tendon sheath+, after which it lies between the heads of the flexor pollicis brevis. It finally attaches onto the base of the distal phalanx of the thumb. It is innervated by the anterior interosseus branch+ of the median nerve+ (C7-C8)

Three dorsal forearm muscles act on the thumb:

The abductor pollicis longus+ (APL) originates on the dorsal sides of both the ulna+ and the radius, and from the interosseous membrane. Passing through the first tendon compartment, it inserts to the base of the first metacarpal bone+. A part of the tendon reaches the trapezium, while another fuses with the tendons of the extensor pollicis brevis and the abductor pollicis brevis. Except for abducting the hand, it flexes the hand towards the palm and abducts it radially. It is innervated by the deep branch of the radial nerve+ (C7-C8).

The extensor pollicis longus+ (EPL) originates on the dorsal side of the ulna and the interosseous membrane. Passing through the third tendon compartment, it is inserted onto the base of the distal phalanx of the thumb. It uses the dorsal tubercle on the lower extremity of the radius as a fulcrum+ to extend the thumb and also dorsiflexes and abducts the hand at the wrist. It is innervated by the deep branch of the radial nerve+ (C7-C8).

The tendons of the extensor pollicis longus and extensor pollicis brevis form what is known as the anatomical snuff box+ (an indentation on the lateral aspect of the thumb at its base) The radial artery can be palpated anteriorly at the wrist(not in the snuffbox).

There are three thenar muscles+:

The abductor pollicis brevis+ (APB) originates on the scaphoid tubercle+ and the flexor retinaculum+. It inserts to the radial sesamoid bone and the proximal phalanx of the thumb. It is innervated by the median nerve+ (C8-T1).

The flexor pollicis brevis+ (FPB) has two heads. The superficial head arises on the flexor retinaculum, while the deep head originates on three carpal bones: the trapezium+, trapezoid+, and capitate+. The muscle is inserted onto the radial sesamoid bone of the metacarpophalangeal joint. It acts to flex, adduct, and abduct the thumb, and is therefore also able to oppose the thumb. The superficial head is innervated by the median nerve+, while the deep head is innervated by the ulnar nerve+ (C8-T1).

Other muscles involved are:

The first dorsal interosseous+, one of the central muscles of the hand, extends from the base of the thumb metacarpal to the radial side of the proximal phalanx of the index finger.

There is a variation in the angle between the first, second phalanges+ of the thumb of humans, varying between 0° and almost 90° when the thumb is extended in a "thumbs-up+". It has been suggested that the variation is an autosomal recessive trait+, called a "Hitchhiker's thumb", with homozygous+ carriers having an angle close to 90°. However this has been disputed, as the variation in thumb angle is continuous and shows little evidence of the bi-modality+ seen in recessive traits.

Malformations of the thumb include a triphalangeal thumb+ and polydactyly+.

One of the earlier significant contributors to the study of hand grips was orthopedic primatologist and paleoanthropologist John Napier+, who proposed organizing the movements of the hand by their anatomical basis as opposed to work done earlier that had only used arbitrary classification. Most of this early work on hand grips had a pragmatic basis as it was intended to narrowly define compensable injuries to the hand, which required an understanding of the anatomical basis of hand movement. Napier proposed two primary prehensile+ grips: the ''precision grip'' and the ''power grip''. The precision and power grip are defined by the position of the thumb and fingers where:
* The '''power grip''' is when the fingers (and sometimes palm) clamp down on an object with the thumb making counter pressure. Examples of the power grip are gripping a hammer, opening a jar using ''both your palm and fingers'', and during pullups.
* The '''precision grip''' is when the intermediate and distal phalanges ("fingertips") and the thumb press against each other. Examples of a precision grip are writing with a pencil, opening a jar ''with the fingertips alone'', and gripping a ball (only if the ball is not tight against the palm).

Opposability of the thumb should not be confused with a precision grip as some animals possess semi-opposable thumbs yet are known to have extensive precision grips (Tufted Capuchins+ for example). Nevertheless, precision grips are usually only found in higher apes, and only in degrees significantly more restricted than in humans.

The pad-to-pad pinch between the thumb and index finger is made possible because of the human ability to passively hyperextend the distal phalanx+ of the index finger. Most non-human primates have to flex their long fingers in order for the small thumb to reach them.

Opposable thumbs are shared by some primate+s, including most catarrhines+. The climbing and suspensory behaviour in orthograde+ apes, such as chimpanzee+s, has resulted in elongated hands while the thumb has remained short. As a result, these primates are unable to perform the pad-to-pad grip associated with opposability. However, in pronograde+ monkeys such as baboon+s, an adaptation to a terrestrial lifestyle has led to reduced digit length and thus hand proportions similar to those of humans. Consequently, these primates have dexterous hands and are able to grasp objects using a pad-to-pad grip. It can thus be difficult to identify hand adaptations to manipulation-related tasks based solely on thumb proportions.

The evolution of the fully opposable thumb is usually associated with ''Homo habilis+'', a forerunner of ''Homo sapiens+''. This, however, is the suggested result of evolution from ''Homo erectus+'' (around 1 mya+) via a series of intermediate anthropoid+ stages, and is therefore a much more complicated link.

It is possible, though, that a more likely scenario may be that the specialized precision gripping hand (equipped with opposable thumb) of ''Homo habilis+'' preceded walking, with the specialized adaptation of the spine, pelvis, and lower extremities preceding a more advanced hand. And, it is logical that a conservative, highly functional adaptation be followed by a series of more complex ones that complement it. With ''Homo habilis+'', an advanced grasping-capable hand was accompanied by facultative bipedalism+, possibly implying, assuming a co-opted evolutionary relationship exists, that the latter resulted from the former as obligate bipedalism was yet to follow. Walking may have been a by-product of busy hands and not vice versa.

HACNS1+ (also known as Human Accelerated Region+ 2) is a gene enhancer+ "that may have contributed to the evolution of the uniquely opposable human thumb, and possibly also modifications in the ankle+ or foot+ that allow humans to walk+ on two legs". Evidence to date shows that of the 110,000 gene enhancer sequences identified in the human genome+, HACNS1 has undergone the most change during the human evolution+ since the chimpanzee-human last common ancestor+.

Many non-human animals also have some kind of opposable thumb or toe. An animal species is said to have ''opposable thumbs'' if the thumb is capable of bending in such a way that it can touch all the other digits on the hand or foot. Most species do not have opposable thumbs. Opposable thumbs are a signature feature of the primate+ family, and played a large role in the ancient humans' invention and use of tool+s.

* Primate+s fall into one of four groups:
** Nonopposable thumbs: tarsier+s and marmoset+s
** Pseudo-opposable thumbs: all strepsirrhine+s and Cebidae+
** Opposable thumbs: Old World monkey+s and all great ape+s
** Opposable with comparatively long thumbs: gibbon+s (or lesser apes)

The thumb is not opposable in all primates — some primates, such as the spider monkey+ and colobus+, are virtually thumbless. The spider monkey compensates for this by using the hairless part of its long, prehensile tail for grabbing objects. In ape+s and Old World monkeys+, the thumbs can be rotated around its axis, but the extensive area of contact between the pulps of the thumb and index finger is a human characteristic.

''Darwinius masillae+'', an Eocene+ primate fossil often described as a missing link between prosimian+ and simian+, had hands and feet with highly flexible digits featuring opposable thumbs and halluces.

* Giant panda+s — five clawed fingers plus an extra-long sesamoid bone+ beside the true first digit that, though not a true digit, works like an opposable thumb.
* In some ''Muridae+'' the hallux is clawless and fully opposable, including arboreal+ species such as ''Hapalomys+'', ''Chiropodomys+'', ''Vandeleuria+'', and ''Chiromyscus+''; and saltatorial+, bipedal species such as ''Notomys+'' and possibly some Gerbillinae+.
* The East African maned rat+ (''Lophiomys imhausi''), an arboreal, porcupine-like rodent, has four digits on its hands and feet and a partially opposable thumb.
Additionally, in many polydactyl cats+, both the innermost and outermost ("pinky+") toes may become opposable, allowing the cat to perform more complex tasks.

* In most phalangerid+ marsupial+s (a family of possum+s) except species ''Trichosurus+'' and ''Wyulda+'' the first and second digits of the forefoot are opposable to the other three. In the hind foot, the first toe is clawless but opposable and provides firm grip on branches. The second and third toes are partly syndactylous+, united by skin at the top joint while the two separate nails serve as hair combs. The fourth and fifth digits are the largest of the hind foot.
* Opossum+s are New World marsupials with opposable thumbs in the hind feet giving these animals their characteristic grasping capability (with the exception of the water opossum+, the webbed feet of which restrict opposability).
* The mouse-like microbiotheres+ were a group of South American marsupials most closely related to Australian marsupials. The only extant member, ''Dromiciops gliroides+'', is not closely related to opossums but has paws similar to these animals, each having opposable toes adapted for gripping.

* The bird-like dinosaur ''Troodon+'' had a partially opposable finger. It is possible that this adaptation was used to better manipulate ground objects or moving undergrowth branches when searching for prey.
* The small, predatory dinosaur ''Bambiraptor+'' may have had mutually opposable first and third fingers and a forelimb maneouverability that would allow the hand to reach its mouth. Its forelimb morphology and range of motion enabled two-handed prehension, one-handed clutching of objects to the chest, and use of the hand as a hook.
* ''Nqwebasaurus+'' — a coelurosaur+ with a long, three-fingered hand which included a partially opposable thumb (a "killer claw").

* Most bird+s have at least one opposable digit on the foot, in various configurations+, but these are seldom called "thumbs". They are more often known simply as hallux+es.

* ''Phyllomedusa+'', a genus of frogs native to South America.

* Prehensility+
* Thumbs signal+

: journal harv
Almécija S. Moyà-Solà S. Alba D. M.
PLoS ONE 5 7 e11727 2010
10.1371/journal.pone.0011727 20661444 2908684

: book harv 2007
Ankel-Simons Friderun
Primate Anatomy
Chapter 8: Postcranial Skeleton
345 3rd
Academic Press 0-12-372576-3

: book harv
Austin Noelle M.
Joint Structure and Function: A Comprehensive Analysis 4th
Chapter 9: The Wrist and Hand Complex
Pamela K. Levangie
Cynthia C. Norkin
F. A. Davis Company Philadelphia
0-8036-1191-9 2005

: book harv
Brown David P.
Freeman Eric D.
Cuccurullo Sara
Freeman Ted L.
Cuccurullo Sara
Physical Medicine and Rehabilitation Board Review
Upper Extremities—Hand Region: Range of Motion of the Digits
Demos Medical Publishing 2004
: book harv 1999
Brunelli Giovanni R.
Brüser Peter
Gilbert Alain
Finger bone and joint injuries Stability in the first carpometacarpal joint
Taylor and Francis 1-85317-690-7

: journal 10.1002/ajp.1350310402 harv 1993
Byrne R.W.
Byrne J.M.E.
American Journal of Primatology 31 4 241–261
John Wiley and Sons

: journal harv
Christel Marianne I.
Kitzel Stefanie
Niemitz Carsten |authorlink3=Carsten Niemitz
How Precisely do Bonobos (Pan paniscus) Grasp Small Objects?
International Journal of Primatology 19 1 165–194
30 November 2004

: journal harv
Costello Michael B.
Fragaszy Dorothy M.
American Journal of Primatology 15 3 235–245
Wiley-Liss March 1988

: journal harv
de Klerk W.J.
Forster C.A.
Sampson S.D.
Chinsamy A.
Ross, C.F.
A new coelurosaurian dinosaur from the Early Cretaceous of South Africa
Journal of Vertebrate Paleontology 2000 2 20 324–332

: journal harv
Diogo R
Richmond BG
Wood B
Journal of Human Evolution 2012 63 1 64–78 | doi=10.1016/j.jhevol.2012.04.001 | pmid=22640954

: book harv
Ellerman John Reeves
British Museum (Natural History) 1941 London

: journal harv
Franzen JL
Gingerich PD
Habersetzer J
Hurum JH
von Koenigswald, W, et al.
PLoS ONE 2009 4 5 e5723
10.1371/journal.pone.0005723 Hawks John

: journal harv
Harcourt-Smith W E H
Aiello L C
Fossils, feet and the evolution of human bipedal locomotion
J Anat
Journal of Anatomy |pmid=15198703 |pmc=1571304 |doi=10.1111/j.0021-8782.2004.00296.x |url=

: journal 10.1016/S0021-9290(08)70148-9 harv
| last=Hsu | first=Ar-Tyan
| coauthors=Meng-Tsu Hu, Fong Ching Su
| title=Effect of Gender, Flexibility and Thumb Type on Thumb Tip Generation
| journal=Journal of Biomechanics | date=July 2008 | volume=41 | issue=Supplement 1 | pages=S148
| url=

: book harv
Jones Lynette A.
Lederman Susan J.
Human hand function
Oxford University Press US 2006

: journal harv
Leakey LSB Louis Leakey
Tobias PV Phillip V. Tobias
Napier JR John Napier (primatologist)
A New Species of Genus Homo from Olduvai Gorge
Nature 4 202 7–9
14166722 | doi=10.1038/202007a0
: book harv 1942
McBride Earl Duwain
Disability evaluation: principles of treatment of compensable injuries
Lippincott 631

: book harv
McDade Melissa C.
Grzimek’s animal life encyclopedia: Volumes 12–16, Mammals I–V 2nd
Koalas (''Phascolartidae'')
Hutchins Michael
Kleiman Devra G.
Geist | editor3-first =Valerius
3 McDade Melissa C.
Farmington Hills, MI Gale Group 2003

: journal 10.1073/pnas.96.1.313 harv
Moyà-Solà Salvador 15136
Köhler Meike 9874815
Rook Lorenzo
PNAS January 5, 1999 96 1 313–317
: journal harv
Napier John Russell
The prehensile movements of the human hand 13376678
J Bone Joint Surg Br 38 4 902–913
November 1956

: book harv
Nowak Ronald M.
Walker's mammals of the world, Volume 2 6th
JHU Press 1999

: book harv 2004
Platzer Werner
Color Atlas of Human Anatomy, Vol. 1: Locomotor System
Thieme 3-13-533305-1 5th

: journal harv
Senter Phil
Comparison of forelimb function between ''Deinonychus'' and ''Bambiraptor'' (Theropoda: Dromaeosauridae)
Journal of Vertebrate Paleontology 2006 26 4 897–906

: journal harv 1946
Slocum D.B.
Pratt D.R.
Disability Evaluation for the Hand
Journal of Bone and Joint Surgery 28 491

: journal harv
van Nierop Onno A.
van der Helm Aadjan
Overbeeke Kees J.
Djajadiningrat Tom J.P.
A natural human hand model
Visual Comput 24 2008 1 31–44

: journal harv |date=January 2003
Young Richard W.
Evolution of the human hand: the role of throwing and clubbing
Journal of Anatomy 202 1 165–174
| pmc=1571064

Human regional anatomy:


Engaging Minds, Transforming Lives

Communication Studies

About the Department

How do we send  messages between one place and another, through some medium of communication? How does the use of different mediums—radio, television, the Internet—affect the way we communicate? How do institutions, such as libraries, maintain and disseminate information over time? These questions, and many others, lie at the heart of the Southwestern Communication Studies Department’s approach.

Through the Department’s multifaceted curriculum, our faculty provides an intimate learning experience, focusing on the myriad ways that communication, culture and identity intersect and define each other.  Students learn a variety of methodologies for the qualitative analysis of communication: ethnographic and historical approaches, as well as approaches located in contemporary feminist theory, critical race theory, queer theory, critical media and cultural studies, and performance studies.

Through this rich combination of modes of thinking and analysis, the Communication Studies Department provides students with a strong critical and theoretical understanding of the ways in which our social reality is constructed and challenged through communication patterns and practices.


The relationship between Lincoln and his military commanders was tested again in January 1864 when the president commuted a death sentence passed on a Union deserter. Unionist generals believed that such a move would undermine discipline in the Union armies. Later in the same month Lincoln did the same thing.

January 1st: The weather greatly hindered nearly all forms of military activity. Sub-zero temperatures occurred as far south as Memphis, Tennessee.

January 2nd: General Banks led his campaign against Galveston by moving along the Texan coast.

January 3rd: Chronic inflation hit the South especially hard. Basic foodstuff was 28 times more expensive in the South than in 1861. During the same time, wages only went up by 3 to 4 times.

January 4th: Lee was given permission by Jefferson Davis to commandeer food in Virginia. Such a measure may have helped his troops but not the people of the state.

January 5th: General Banks was encouraged by General Halleck to be more aggressive during his offensive. Halleck envisaged Union troops in Galveston by the spring.

January 7th: Lincoln commuted the death sentence imposed on a Union deserter. His move, as commander-in-chief, was not well received by the Union’s military hierarchy who felt that it would undermine discipline even more. Union desertion was at an all-time high, especially in the Army of the Potomac. The problems were many but one of them was that men were paid a bounty to enlist. Many did then deserted only to enlist again to collect another bounty. Also if you name came up in a draft for your town/city you could exempt yourself by paying $300. Or you could get a substitute to do your draft for you. Big cities saw a major growth area in “substitute brokers” who, for a fee, would find a substitute for someone. The whole system was open to abuse and when 57 men were recruited to the 6th New York Heavy Artillery, 17 were physically disabled.

January 8th: David O Dodd, convicted of being a Confederate spy, was hanged in Little Rock, Arkansas.

January 10th: The Confederacy responded to its economic plight by printing more money. Foreign governments were unwilling to lend money to it and only accepted gold for the payment of weapons. The North made matters worse for the South by printing counterfeit Confederate notes, which made confusion endemic.

January 11th: Senator John Henderson (Missouri) proposed within the Senate that slavery should be abolished throughout the USA.

January 18th: Protest meetings were held in North Carolina regarding the conscription policy of the Confederacy. All white males between 18 and 45 were required to enlist – shortly to increase to all males between 17 and 60.

January 19th: Pro-Union supporters met at Little Rock, Arkansas.

January 21st: Pro-Union supporters met in Nashville, Tennessee.

January 23rd: Lincoln approved a plan that allowed plantation owners to hire those who had previously worked as slaves on their plantations.

January 26th: Lincoln commuted another 9 planned executions, as he did not want to “add to the butchering business”. On the same day he approved a plan to improve trade between the Union and those parts of the Confederacy now under Union control. Lincoln was already thinking beyond the end of the war and wanted to ‘normalise’ internal trade as much as possible.

January 31st: Lincoln stated that he hoped all former slaves who wanted to fight for the Union would swear the oath but that it was not an absolute requirement. The same was true for those men who had been in Confederate ranks – swearing loyalty to the Union was preferred but was not absolute.


Economist says Sacramento Kings fight may yield antitrust lawsuit

Apr 8, 2013

"Whichever city is going to lose has an antitrust case against the NBA for not having a team in that city," Noll said.

"The crucial issue is whether the purpose and effect of the restriction is simply to enhance the monopoly power of the existing set of teams," Noll said.

He says professional sports leagues have manage to duck this issue by expanding their leagues whenever the issue arose in the past. And he expects the same thing will happen this time. Both Seattle and Sacramento will eventually have their own teams.


You are here

Fires in the Bathroom: Advice for Teachers from High School Students

Kathleen Cushman
The New Press
Publication Date:
Number of Pages:
[Reviewed by
Kara Shane Colley
, on

Fires in the Bathroom: Advice for Teachers from High School Students might seem like a scary title for a teacher to read.  What awful things might these teenagers reveal about my teaching?  Fear not.  Kathleen Cushman and the student co-authors from What Kids Can Do have assembled thoughtful and respectful advice on things their beloved teachers do right and mistakes even the best teachers make.  Fires in the Bathroom is not so much about curricular suggestions; it focuses instead on the "relationships that make learning possible."  The book includes advice on behavior management, gaining the trust and respect of students, motivating students to learn, and teaching English language learners.  For example, Alexis gives the book's best classroom management advice, "When you let your students walk all over you, they're not learning the essentials of respect, of how to interact properly."  The authors also have some interesting tips, like homework do's and don'ts, possible punishments, and timing of tests.  Overall, what comes through loud and clear is that students want authoritative, yet respectful teachers, and that they want to learn, to succeed, and to feel special yet blend in.

My favorite part of the book is the worksheets for teachers.  There is one on "Identifying the Assets of your Worst Behaving Students."  It aims for the teacher to look at other aspects of the student's life in order to gain insight into the disruptive behavior.  Another good one is the worksheet called "I Don't Get It" which helps teachers remember the experience of being confused in order to understand what students may be going through.  The worksheets are helpful little homework assignments for teachers.

Fires in the Bathroom does raise two questions.  The whole premise of the book is that students have important ideas about their schooling.  This makes me wonder how much control students should have over their education.  Should they have input about what to learn, when to learn it, and how to learn it?  Second, there is a small section in the book on how to teach math.  Most of the advice is typical, such as "Use lots of concrete examples" and "Try all different ways to see something."  One piece of advice is "Walk us through things."  I find this particularly interesting in light of the recent emphasis in math education on problem solving.  Alexis writes "He doesn't want to tell us how to do it — he wants us to figure out for ourselves.  It's hard.  It's frustrating.  It feels like he's not teaching."  What is the balance between leading students through a problem and letting them wrestle with it?  It is a tricky dilemma that math educators need to explore further.

Fires in the Bathroom would make excellent reading for a first year teacher.  It is a quick read and would fit well in a teacher preparation program.  Any teacher that cares about her classroom relationships could use this book as a starting point for dialogue with her students.  A teacher who is feeling discouraged might take heart when advised not to "judge your success by whether students like you."  Vance puts it best, "You really affect kids when you just do your job, day in and day out, do it well — and everything doesn't have to be about bonding with the kids and changing their lives.  That's artificial.  The bond will develop on its own if you just do your job well."

The table of contents is not available.


Marine comes home to big surprise

After more than two decades in the Service, most recently in Okinawa, a Northern California Marine was ready to settle down. So, the retired Marine Master Sgt. Jacinto Bernardo bought a house -- but it needed a lot of repairs. Here's what happened when he pulled up to his driveway Tuesday. Surprise!


What does WPWL stand for?

What does WPWL mean? This page is about the various possible meanings of the acronym, abbreviation, shorthand or slang term: WPWL.

We've found a total of 1 definition for WPWL:

Filter by: Sort by:PopularityAlphabeticallyCategory

Water Polo World League

Rate it:

Know what is WPWL? Got another good explanation for WPWL? Don't keep it to yourself!

"WPWL." Abbreviations.com. STANDS4 LLC, 2016. Web. 14 Feb. 2016. <http://www.abbreviations.com/WPWL>.


People urged to avoid swimming in Lake Tutira

Hawke’s Bay DHB Medical Officer of Health Dr Nicholas Jones says that the mostly likely cause of rash occurring among swimmers at Lake Tutira is “swimmers itch”.  Also known sometimes as “duck itch” the rash is due an allergic reaction to bird parasites.  Dr Jones said he is not aware of swimmers itch occurring previously in Hawke’s Bay but it has been known to cause problems in other parts of the country such as Lake Wanaka.

It is possible that only some areas of the lake are affected.

Dr Jones advice at this time is to avoid swimming in Lake Tutira. If people do choose to swim it would be particularly important to avoid swimming in the shallow waters at the lake edge.  This makes it particularly difficult for parents with young children so parents may decide it is safer to keep children out of the water altogether or anywhere near the lake edge.

People who are experiencing itch or rash after swimming at the lake should see their family doctor for assessment and treatment.  To discuss other concerns or report where the rash occurred please contact the DHB’s Public Health Unit available through 06 878 8109.

Dr Jones said signs were erected on Thursday at the lake to warn visitors and the DHB was working with the Regional Council to further assess both immediate and longer term risks.

How does “swimmers itch” occur?
Swimmers itch or duck itch is a rash due an allergic reaction to bird parasites The parasites have a life cycle that relies on growth in snails before re-infecting aquatic birds, the snails tend to live in the shallow water around the lake edge
If someone is unfortunate enough to swim at a location where snails are releasing the parasite the rash is likely to occur as the parasite enters the swimmers skin.  Although the parasite does not survive in humans the rash occurs because of the body’s immune system reaction.
Swimmers may initially experience itch or tingling on leaving the water which develops into an intense itch over a number of hours as red spots enlarge to form pimple like bumps.  The rash usually only occurs in areas of the body that have been exposed to the affected water.

How is “swimmers itch” treated?
When there are only a few spots no treatment may be necessary. Otherwise the skin should be lightly rinsed with (rubbing alcohol) and coated in calamine lotion.  Antihistamine or mild corticosteroid cream can be helpful.
For some forms of the parasite brisk towelling immediately after swimming can reduce the risk of a reaction developing. However at this stage not enough is known about the cause to determine whether towelling would be helpful in this instance.

More information is available at


Fuel Cells

NYSERDA's Fuel Cell programs support the installation of fuel cells for businesses, hospitals, multifamily buildings, and other large power consumers.

Fuel cells convert natural gas to a hydrogen-rich fuel that is combined with atmospheric oxygen in an electrochemical process to produce electricity, heat, and water. Fuel cells emit significantly less carbon dioxide than conventional combustion engines and give off virtually none of the harmful pollutants associated with burning fossil fuels.

Financial incentives are available to support the installation and operation of continuous duty fuel cell systems in New York State through the Renewable Portfolio Standard (RPS) Customer-Sited Tier - PON 2157 and the RPS Main Tier


The state has never regulated "fracking" and the oil and gas industry has alternatives, Longmont attorneys argued in a brief late Friday defending its ban of hydraulic fracturing.

The brief asks the Boulder County District Court to not summarily kill the ban, but to let it go on to an evidentiary hearing in April. The Colorado Oil and Gas Association, which sued Longmont over the ban in 2012, asked for a summary judgment in March.

Longmont residents voted in 2012 to ban fracking, which uses high pressure fluid to break open hard-to-reach oil and gas deposits. The ban, which was adopted into the city charter, also forbids the storage of fracking wastes in city limits.

COGA has maintained that the ban is tantamount to a ban on drilling — forbidden in 1992 by a Colorado Supreme Court decision — and that only the state can regulate the methods used.

But the state hasn't done that in this case, argued city attorney Eugene Mei, assistant city attorney Dan Kramer and special counsel Phillip Barber. Hydraulic fracturing was never written into the state law that empowered the COGCC, they wrote, and the COGCC's own rules merely call for operators to give a 48-hour warning of a frack and report any chemicals used.

"(T)he COGCC's rules amount to non-regulation of hydraulic fracturing," the city's lawyers wrote. "The COGCC does not issue permits for it. The drilling permit from the COGCC does not 'tell the operator yes, you are allowed to hydraulically fracture, or, no, you're not allowed'; it's up to the operator to make that decision."

By contrast, they noted, there are COGCC rules that regulate several other aspects of drilling, down to locating signs and cleaning up trash.

"The absence of any comparable rules for fracking speaks volumes," the attorneys wrote. There can't be an "operational conflict" between state and local law, they argued, if there are no state regulations to conflict with.

"The COGCC. . . has abdicated its oversight responsibility to the oil companies," the brief said.

The brief also argues that fracking presents health hazards to humans and animals; risks of spills, air pollution and earthquakes; increased traffic; and the need for local fire departments to get additional equipment and training for well fires.

The local interests involved are huge, the brief stated, and the ban affects only a "minuscule" portion of the oil and gas to be had in Colorado.

"For that small patch of land," Mei, Kramer and Barber wrote, "the citizens of Longmont are achieving the state interest better than the state itself: to allow production while protecting human health, safety and the environment."

The first test of the ban came in 2013 when Synergy gained permits for 23 wells in Firestone's Union annexation near the Longmont city limits. The company drilled the wells but chose not to frack the areas that passed beneath Longmont's bounds rather than get into a legal battle of its own.

Far from being damaged, Longmont's attorneys argued, Synergy's profits had gone up 89 percent in the last year.

The brief lists wells in the Wattenberg Field — the large oil and gas field that Longmont is on the edge of — that were completed without fracking. It also mentions an alternative method called "underbalanced drilling" or UBD that uses the natural pressure of the well to bring up oil and gas without the need to frack.

"Economic wells have been drilled without hydraulic fracturing, including wells drilled with the UBD technique," Longmont's attorneys wrote. "Likewise, there are many wells that have been completed using hydraulic fracturing that become uneconomic in a few months. Thus, hydraulic fracturing is not the only completion technology which would result in a producing or economic well."

COGA's request for summary judgment has been echoed by the Colorado Oil and Gas Conservation Commission — the state regulating body for the industry — and the drilling company TOP Operating.

The COGCC has a separate lawsuit with Longmont over drilling regulations the city adopted in 2012, including a restriction against drilling in residential zones. Those rules did not include a fracking ban. The case was originally set to be heard this fall, but was postponed so that the suit against the fracking ban could be heard first.

The case is being heard before Judge D.D. Mallard.

Summary judgments can be granted if there are no factual issues that would require a trial, and if it can be shown that one side or the other clearly has the law on their side.


Discover and save creative ideas
    Explore these ideas and more!

    Bioluminescent Phytoplankton / Vaadhoo Island, Maldives

    Beach in the Maldive islands that glows with millions of pinpoints of glowing blue. The light from these bioluminescent phytoplankton looks like a fantastic starry sky somewhere deep in the universe.

    bioluminescence on a Maldive beach

    Hidden beach on Marieta Islands, Mexico

    Melissani Cave @ Kefalonia, Greece

    101 Most Beautiful Places To Visit Before You Die!

    cherry blossom lake @ sakura japan

    Castle Island, Dublin

    Those glowing blue blobs are actually deep sea creatures called Glowing Firefly Squids, and they are some of the most beautiful of earth's marine life.

    Blue Caves, Greece

    Capri - Campania, Italy

    Hillier Lake, Western Australia: The pink and lovely Hiller Lake is the only vividly pink lake you will find in the world. The color is permanent and never changes, even when water is removed and placed in a separate container. Its startling color remains a mystery and while scientists have proven it's not due to the presence of algae, unlike the other salt lakes down under, they still can't explain why it's pink....

    Sunset at Kiholo Bay, Hawaii

    Conrad Maldives Rangali Island(10+ Pics)

    The Holy Hole, Madeira Island, Portugal

    Huka Falls, New Zealand

    41 Places To See Before You Die

    Maldives (25+ Pictures)

    Socotra Island, Yemen


Sunday, February 14, 2016
Gateway to Sikhism

The Indian Mutiny - 1857

The 14th King Georges own Ferozepore Sikhs.

At this time the Army in India was organized in three Presidency armies of Madras, Bombay and Bengal. Each army consisted of units of the British Regular Army, known as "Royal" troops, and British and Indian units of the East India Company. The Bengal Presidency stretched right across India from Peshawar to Calcutta, which was the headquarters of the British Government. The majority of the British troops of the Bengal Army were stationed in the Punjab, which had only recently been acquired. Although the importance of Allahabad was recognized by the Government, there were no effective British troops stationed in the fort and there were no British reserves between Calcutta and Delhi.

The following is an extract from the London Times of that time

On the same day General Havelock arrived in Allahabad with the 64th and 84th Foot and the 78th Highlanders, and he set off for Cawnpore a few days later, taking with him his British troops and a detachment of the Ferozepore Sikhs. By this time Cawnpore had been captured by the rebels, so General Havelock decided to drive them out and then march to the relief of Lucknow, where the British were besieged in the Residency.

A portion of the Ferozepore Sikhs were left behind in Allahabad, under Lieutenant Montague, to hold the fort and patrol the surrounding district. Here the Sikhs did excellent work and fought several successful engagements with parties of mutineers in the area. On one occasion a guard of two non-commissioned officers and eight sepoys, surrounded by about a thousand rebels at Sahunga, gallantly rescued a wounded British officer and fought their way back through the rebels to the main guard.

General Havelock joined forces with the advanced column on the 12th of July and moved on towards Cawnpore in very trying conditions in the hot weather. On the following day, just as the combined force was preparing to camp

near the village of Fathepur, a large party of mutineers advanced from the village to attack the British force. Although his men were exhausted after a long march under a scorching sun, Havelock decided to attack. He immediately deployed his troops and utterly routed the enemy in a short, sharp fight. After a much-needed rest on the next day, the force continued the march early on the 15th of July. However, it was found that the enemy had re-formed and was holding the village of Aong in strength. General Havelock immediately attacked the enemy positions and threw back the mutineers at the point of the bayonet. It was now learnt that the enemy was preparing to blow the important bridge over the Pandu river, six miles farther on, so Havelock had to push on without resting in order to save the bridge and secure a passage over the river. Brasyer's Sikhs moved forward in skirmishing order and occupied the cliffs overlooking the bridge. This enabled the guns to come forward and cover the Madras Fusiliers, who stormed the bridge and put the enemy to flight.

The same evening General Havelock learnt that a number of women and children had been made prisoner at Cawnpore and had to be rescued at all costs. He therefore decided to continue the advance without delay, even though his men had had no rest and the column was still twenty-two miles from Cawnpore On the 16th of July the force advanced to within a few miles of the town before meeting any resistance. Here some ten thousand rebels opposed the British advance on the town. General Havelock personally led his now-small force of nine hundred men round the enemy's left flank and took the enemy by surprise from the rear. The 78th Highlanders were in the lead and rolled up the enemy's left flank with a brilliant charge. The 64th and 84th Foot and Brasyer's Sikhs then passed through and carried the enemy's position. They captured the guns on the right and the enemy retreated. Leaving the guns behind, protected by Brasyer's Sikhs, the British infantry regiments followed up their success and inflicted further losses on the enemy, who eventually lost heart and fled in disorder.

General Havelock and his men camped for the night in the open and entered Cawnpore early on the 17th of July, but they were too late to stop the brutal murder of the women and children by the mutineers.
Forest, in his "History of the Indian Mutiny," wrote as follows about
Havelock's advance from Allahabad
" In nine days Havelock and his veterans had marched 126 miles under an Indian sun in the hottest season of the year, each man carrying a heavy weight of ammunition, and had won four pitched battles and sundry combats against highly disciplined troops far exceeding them in number. During four days' fighting they had killed or wounded many hundreds of their enemies, and had captured twenty-three pieces of artillery. Their advance had been one of suffering, of privation, and of fatigue. . . . Battle after battle was won by desperate fighting; the cholera and the sunstroke slew many survivors of the combat, but on they went with unflinching resolution until Cawnpore was reached."

After a few days' rest Havelock, leaving General Neill with a small force to hold Cawnpore, crossed the River Ganges by boat and set out to march to the relief of Lucknow, forty-five miles away. His force, which was only fifteen hundred strong and included Brasyer's Sikhs, moved out on the 29th of July and almost immediately encountered a large force of the enemy opposing their advance. Havelock drove the enemy out of the villages of Unao and Basiratganj and utterly defeated them in two brilliant battles. However, Havelock's force was seriously depleted by sickness and battle casualties and he had to withdraw to Mangalwar; a few miles north of the river, and await reinforcements. It was quite obvious that the remnants of his force had little chance of forcing the way to Lucknow and carrying out the relief of the besieged garrison in the Residency. Forrest wrote in his History
" Two victories had been won. But if the road to Lucknow was to be so roughly contested there was little chance of reaching the Residency. What soldiers could do Havelock's men had achieved. But they could not fight the pestilence of the tropics. For some days cholera and dysentery had done deadly work among them. A sixth of his force had perished-half on the battlefield, half by disease."

A few days later Havelock received a small number of reinforcements and a few guns, so he moved forward again on the 5th of August. He encountered the enemy in Basiratganj and utterly routed the rebels for a second time, but again was forced to withdraw to Mangalwar. He was still not strong enough to fight his way to Lucknow, which was reported to be held by thirty thousand mutineers.

On the 11th of August Cawnpore was threatened by four thousand mutineers, who had arrived in Bithur from Saugor, and General Neill called for aid, while, at the same time, the enemy was also reported to be collecting again in Basiratganj. Havelock was determined to strike another blow before recrossing the river to Cawnpore, and he set out with his force the same evening. He once again defeated the enemy in a fierce battle a few miles north of Basiratganj during the next morning, and then withdrew for a third time and crossed the river to Cawnpore.

Statue of Havelock in Trafalgar Square

In the middle of September Sir James Outram arrived in Cawnpore with a large force of reinforcements and bridging operations over the Ganges were begun. The mutineers attacked the bridge from the northern bank and Brasyer's Sikhs were sent over to cover the construction. The Sikhs drove the enemy back and the bridge was completed without further interference.

On the 21st of September two brigades, about three thousand strong all told, set out for Lucknow under General Havelock, accompanied by Sir James Outram.

The enemy opposed the advance at Mangalwar and at Alambagh, in the southern outskirts of Lucknow, and were utterly defeated by the British in two gallant battles. Havelock and Outram halted at Alambagh on the 24th of September while they decided the best means of extricating the British forces in the Residency.

The sick and wounded, heavy baggage and large supply train were left at Alambagh, protected by a guard of three hundred men drawn from all units, in the force.

The rearguard, with a number of sick and wounded, had not been able to reach the Residency and had remained in the Moti Mahal. So, on the next day, a detachment of the 5th Fusiliers and Brasyer's Sikhs was sent to reinforce them and help them to withdraw to the Residency. Although the Sikhs and Fusiliers fought their way through and drove the enemy back from the buildings and gardens adjacent to the Mod Mahal, the enemy fire from the Kaiserbagh was found to be too heavy to admit of the rearguard convoy being moved back. Further reinforcements from the 78th Highlanders were then sent forward and the rearguard was safely withdrawn to the Residency after dark.

After arriving in the Residency area Sir James Outram took over, from General Havelock, the command of the British forces. Although the rebels had been outwitted, they had not been decisively defeated and still occupied the city in great strength. It was found to be quite impracticable to carry out the original intention of withdrawing the besieged people in the Residency and all the relieving force could do was to aid its defences. Although this was not really a relief of the Residency, it was a very gallant rescue from a situation of the gravest peril. There were now 2,000 additional troops, so there was no longer an imminent danger of the garrison being overwhelmed. However, the Residency was besieged as closely as ever, and Sir James Outram had to stand on the defensive and await relief in his turn.

With the increased number of troops in the Residency positions had to be enlarged and so for the next few days several sorties were made to improve the position. The Regiment of Ferozepore was in General Havelock's sector and took part in the sorties along the eastern face of the Residency to clear the enemy from the gardens and houses up to the Chata Manzil. These sorties were entirely successful and improved the defences of the Residency. Lieutenant Cross, of the Ferozepore Sikhs, was wounded in one of these sorties, but otherwise the Regiment suffered very few casualties.

At last, on the 17th of November, a relieving force under General Sir Colin Campbell, Commander-in-Chief in India, arrived at Lucknow. The situation at Cawnpore, however, had again become: critical and General Campbell had to return there as quickly as possible. He therefore decided to evacuate the Residency and return to deal with the rebels at Lucknow at a later date. On the night of the 22nd November all the British forces were withdrawn successfully from the Residency together with all the women, children and wounded. The enemy were taken completely by surprise by this operation, which had been carefully planned and boldly executed.

General Outram was left with a force of some four thousand men to hold Alambagh and contain the enemy at Lucknow. The Ferozepore Regiment was included in General Outram's force and held defensive works at Alambagh for three months. Duties were very arduous on account of the large perimeter to be held. while the enemy kept in constant touch and there were almost daily skirmishes and minor encounters. The enemy delivered a number of attacks, but these were all beaten off with losses to the rebels.

On the 22nd of December General Outram took the offensive and threw back a large enemy force which had attempted to sever his communications to Cawnpore. Reporting on this action, Outram wrote
" The gallant way in which, with a, cheer, the 78th and the Regiment of Ferozepore, led by their commanders, dashed at a strong position held by the enemy (30,000 men and 6 heavy guns), excited much admiration."

At the beginning of March, 1858, Sir Colin Campbell, with a large, well equipped force, joined General Outram at Alambagh and started methodical operations against the rebels at Lucknow,, The enemy were holding three lines of defences north of the city covering the Kaiserbagh, their citadel. These had been strengthened since the relief of the Residency, and houses were now fortified and roads barricaded.
Sir Colin's plan was to send General Outram with his division north of the River Gumti to turn the rebels' position, while his main force attacked the Kaiserbagh from Dilkusha Park.

General Franks, in his report of that day, wrote


Dr Daniel Sadigh

A Moment with Self

Follow This Show

Stay in the know about new episodes and updates.
Covers your relationship with your self as well as with others. Dr. Daniel Sadigh discusses subjects such as family issues, stress, anxiety, depression, and financial success.

On-Demand Episodes

Effective communication is the most important element in every healthy relationship. Personal sensitivities can trigger emotional reactions and easily derail our connections to others. If we are not able to accurately express... more

Have you ever found yourself talking to yourself?How we feel about a situation and what we do with it in action requires paying attention to an internal dialogue. However, we are often confused by the prevalent language in our heads as the... more

Ending a relationship has never been easy. The physical and emotional suffering of a breakup and the confused state that follows can become torturous. How can one start over and fall in love again after experiencing the pains of a... more

If you have ever wondered what homeopathy is and how it is perceived to alleviate suffering, this interview is your opportunity. Based on the doctrine of "like cures like" a substance that causes the symptoms of a disease in healthy... more

Have you ever wondered why some people can behave unemphatic, arrogant, envious, entitled, and shameless in relation with their own family members? Have you noticed how these people overestimate their abilities and have excessive... more

The emotional, social, and practical challenges for women without children, by circumstance, is often an unspoken subject. According to Marcy Cole, Ph.D"There is a mother that lies within the heart of every woman?.Dr. Cole is a Holistic... more

Do you ever doubt your decisions? Do you procrastinate? Do you believe you deserve negative outcomes? How about feeling able to overcome challenges? Familial traits, socio-economic backgrounds, abusive relationships, levels of... more

What is bullying and why do bullies bully? Aggressive or dominating behavior is often displayed by teenagers and can continue over a lifetime in abusive relationships. Such a primitive psychological defense mechanism finds roots in... more

The knowledge and understanding of what an elder feels during the Holidays can help families make better plans. There are many questions to be asked -- and answered -- about the treatment of elders and their inclusion in joyous, often... more