text:
A whole genome Bayesian scan for adaptive genetic divergence in West African cattle
© Gautier et al; licensee BioMed Central Ltd. 2009
Received: 29 May 2009
Accepted: 21 November 2009
Published: 21 November 2009
The recent settlement of cattle in West Africa after several waves of migration from remote centres of domestication has imposed dramatic changes in their environmental conditions, in particular through exposure to new pathogens. West African cattle populations thus represent an appealing model to unravel the genome response to adaptation to tropical conditions. The purpose of this study was to identify footprints of adaptive selection at the whole genome level in a newly collected data set comprising 36,320 SNPs genotyped in 9 West African cattle populations.
After a detailed analysis of population structure, we performed a scan for SNP differentiation via a previously proposed Bayesian procedure including extensions to improve the detection of loci under selection. Based on these results we identified 53 genomic regions and 42 strong candidate genes. Their physiological functions were mainly related to immune response (MHC region which was found under strong balancing selection, CD79A, CXCR4, DLK1, RFX3, SEMA4A, TICAM1 and TRIM21), nervous system (NEUROD6, OLFM2, MAGI1, SEMA4A and HTR4) and skin and hair properties (EDNRB, TRSP1 and KRTAP8-1).
The main possible underlying selective pressures may be related to climatic conditions but also to the host response to pathogens such as Trypanosoma(sp). Overall, these results might open the way towards the identification of important variants involved in adaptation to tropical conditions and in particular to resistance to tropical infectious diseases.
Cattle are still playing a major role in Africa for food supply, to generate income and draught power or for ceremonial purposes. Archaeological, historical and anthropological evidence combined with recent genetic data [1] have provided insights into the complex origins of present day West-African cattle diversity. Indeed, although their wild ancestor Bos primigenius was not native to sub-Saharan Africa, West African cattle populations are representative of both shorthorn (Bos taurus brachyceros) and longhorn (Bos taurus longifrons) humpless taurines, humped zebus (Bos indicus) and zebu/taurine hybrid cattle. This early suggested that West African cattle has originated from several successive and recent colonization events [2, 3]. Briefly, shorthorn taurines were introduced from the Middle-East and possibly North Africa around 4,000 years BP [3, 4] while longhorn taurine probably arrived at an earlier period (5,000 years BP) following different migration routes [3]. Although, zebu cattle first penetrated through the Horn of Africa in the late 2nd millennium BC, the major wave of indicine introgression really started with the Arab settlements along the East Coast of Africa from the end of the 7th century AD. Zebu cattle spread even more recently over West Africa with movements of pastoralist people such as the Fulani [1].
West African cattle populations thus represent an appealing model to unravel the genome response to adaptation to tropical conditions. The purpose of this study was to perform a whole genome scan for footprints of adaptive selection based on a newly collected genotyping data set containing 36,320 SNPs genotyped on 9 West African cattle populations from different bovine sub-species and agro-ecological areas (Figure 1). In particular, we sampled populations on both side of the tsetse infested zone. Based on this large data set, we first carried out a detailed analysis of the genetic structure of these populations. We next performed a scan for differentiation among SNPs under a full Bayesian framework [7, 8] for which we proposed additional extensions to improve the detection of loci under selection. We then annotated regions containing SNPs subjected to selection, adopting a systems biology approach to highlight the main underlying physiological functions.
Results and Discussion
SNP polymorphism
In total, 437 individuals belonging to 12 different cattle populations, nine of which originating from West Africa (Figure 1), were genotyped for the BovineSNP50 chip assay containing 54,001 SNPs mainly derived from sequences available in European cattle breeds [9]. Among the autosomal SNPs that passed Quality Control analyses (see Methods), we retained the 36,320 SNPs polymorphic (Minor Allele Frequency or MAF above 0.01) in at least one West-African taurine and one West African zebu populations. As expected and shown in Figure S1A (additional file 1), this SNP selection procedure, leading mainly to the elimination of SNPs from European origin, resulted in a relative increase in calculated heterozygosity for all but outgroup taurine populations (Aubrac or AUB and Oulmès Zaer or OUL). In addition, as revealed by the distribution of the MAF for the different populations (Figure S1B in additional file 1), the remaining differences in heterozygosity among populations were mainly explained by the proportion of SNPs with rare variants (MAF < 0.05). In particular, marker polymorphism remained clearly lower in West African taurine populations (ND1, ND2, LAG, SOM and BAO), the proportion of SNPs with a MAF < 0.05 being the highest (36%) in LAG which also displayed the lowest average heterozygosity (0.118). These observations are in agreement with previous studies based on microsatellite markers on the same populations [10] and might essentially be explained by the recent demographic history of West African taurine cattle characterized by a marked isolation associated with a strong effective population size decrease (e.g. [11]). Although West African zebus (ZFU and ZCH) and hybrids (BOR and KUR) displayed more genetic diversity than West African taurines in our study (Figure S1 in additional file 1), ZFU, ZCH and ZMA levels of polymorphism were clearly lower than, and BOR and KUR ones similar to the taurine outgroup (AUB and OUL) ones (Figure S1A in additional file 1). Nevertheless, studies based on microsatellite data (e.g. [10]) revealed that heterozygosities of ZCH, ZFU, BOR and KUR were higher than European breeds' ones (AUB) which were themselves similar to ZMA. The lack of zebu specific sequences in the SNP discovery process might explain these apparent discrepancies.
Analysis of Population Structure
As reported in Table S1 (additional file 2), within population F IS were almost null (< 0.006) for ZFU and BOR or very close to zero (from -0.0121 to 0.0198) for ND1, BAO, KUR and ZCH. The slightly positive F IS (0.0421) observed in SOM might result from the sampling area being apart the border between Togo and Benin. The positive F IS (0.0414) of similar magnitude observed in LAG might rather originate from a higher level of inbreeding as suggested by the extent of Linkage Disequilibrium (LD) within this population (see below). Conversely, for ND2, the negative F IS (-0.108) might be explained by recent outcrossing events. Indeed ND2 individuals derived from founders recently collected in different and distant Ivorian villages (see Methods). This hypothesis is also in agreement with their higher dispersion compared to other West African taurines in the PCA (Figure 2) and a higher level of overall extent of LD (see below). Nevertheless, the global F IS coefficient remained close to zero (0.0103) while the global F ST and F IT were respectively equal to 0.132 and 0.141. Such a level of differentiation among the West African populations was in agreement with the one computed based on microsatellite markers [10] suggesting a low impact of the SNP ascertainment procedure on these estimates.
Bayesian scan for differentiation among SNPs
We further studied the differentiation among SNPs and populations based on a Bayesian hierarchical model derived from [7] and presented in more details in the Methods section. Among the factors influencing population differences in allele counts (differentiation), the model aims at distinguishing locus-specific from population-specific factors, such as migration or drift [17]. Because most SNPs originated prior to breed differentiation, the effect of different mutation rates across loci might be negligible and the SNP-specific effect in the model is mainly related to selection. Notice that the model assumed the F IS to be null within each population which appeared sound given the characteristics of our data (see Figure S3 in additional file 1 and Methods). We also ignored ascertainment biases originating both from the SNP discovery process and our selection of SNPs "informative" in West African populations. These two different sources are expected to favour polymorphic SNPs of ancient origin (co-segregating in European and African cattle breeds) and thus modifies to some extent the distribution of allele frequencies in the gene pool (x following our model notations) [18]. Recently, efficient algorithms were proposed to account for ascertainment biases [19], providing it can be modelled (which is difficult in our case at least for the first identified source). Nevertheless, for the purpose of this study we were mainly concerned with possible biases introduced in the locus-specific effect estimation which might not be of importance [18]. At least for our second source of ascertainment bias, simulated data allowed to confirm this statement (Figure S4 in additional file 1) since estimation of x was found robust to departure from the prior distribution assumed. Similarly, although more variable, locus-specific effect estimates were highly correlated with their corresponding simulated values (r > 0.8).
FDR (FNR) for different thresholds on BF.
BF threshold (in dB units)
1.40 (70.0)
2.98 (56)
14.0 (37.9)
28.5 (27.6)
0.477 (78.0)
1.07 (66.2)
4.75 (50.1)
8.84 (44.4)
0.166 (84.0)
0.364 (75.4)
1.71 (61.1)
3.23 (55.7)
0.0501 (88.0)
0.158 (82.6)
0.653 (71.3)
1.23 (66.1)
0.0180 (88.0)
0.0788 (86.2)
0.231 (79.9)
0.418 (76.2)
0.0120 (88.0)
0.0384 (87.8)
0.100 (84.3)
0.172 (82.1)
0.0120 (88.0)
0.0242 (88.4)
0.0578 (86.9)
0.0934 (85.4)
0.0120 (90.0)
0.0242 (89.4)
0.0400 (88.7)
0.0369 (87.3)
0.00 (100)
0.0182 (89.6)
0.0333 (89.5)
0.0197 (88.1)
0.00 (100)
0.0141 (90.0)
0.0333 (89.8)
0.0172 (88.5)
0.00 (100)
0.0141 (90.0)
0.0311 (89.9)
0.0172 (88.7)
Relationship between selection coefficient (s) and locus effect α i
Simulated s
Number of loci simulated
Average FST (sd)
FST range (median)
α i
0.128 (0.06)
0.007-0.479 (0.121)
0.173 (0.085)
0.019-0.479 (0.166)
0.189 (0.105)
0.012-0.646 (0.171)
0.21 (0.108)
0.025-0.56 (0.196)
0.223 (0.113)
0.024-0.585 (0.211)
0.327 (0.152)
0.019-0.729 (0.348)
0.439 (0.183)
0.041-0.748 (0.489)
0.509 (0.194)
0.068-0.841 (0.572)
0.527 (0.199)
0.026-0.838 (0.588)
0.57 (0.242)
0.018-0.887 (0.677)
0.649 (0.227)
0.068-0.925 (0.728)
0.71 (0.239)
0.04-0.972 (0.801)
0.12 (0.052)
0.006-0.339 (0.113)
0.11 (0.049)
0.017-0.305 (0.104)
0.104 (0.047)
0.014-0.269 (0.096)
0.091 (0.044)
0.002-0.267 (0.086)
0.07 (0.039)
0.005-0.286 (0.061)
0.056 (0.036)
0.006-0.238 (0.048)
0.051 (0.037)
0.002-0.177 (0.043)
0.05 (0.034)
0.004-0.203 (0.042)
0.041 (0.033)
0.001-0.19 (0.031)
0.04 (0.034)
0.001-0.22 (0.029)
0.035 (0.034)
0-0.187 (0.027)
A Genome map of regions under adaptive differentiation
Regions under selection identified at the 5% local FDR (q-value) thresholds.
Position (size) in Mb
Peak Position in Mb
BF value of the SNP at the peak position (q-value of the smoothed signal)
Gene at or close to the peak
3.654-12.74 (9.087)
50 (0.0467)
133.1-138.6 (5.465)
51 (0.0345)
0.035-2.687 (2.652)
51 (0.0001)
64.04-67.28 (3.235)
31 (0.0258)
71.48-78.32 (6.84)
51 (0.0001)
117.7-129.0 (11.35)
45 (0.0007)
140.0-140.6 (0.662)
25 (0.0473)
12.39-19.72 (7.325)
51 (0.0007)
107.9-110.1 (2.219)
37 (0.0246)
24.74-29.70 (4.953)
38 (0.0002)
32.31-37.17 (4.860)
36 (0.0009)
46.84-54.10 (7.259)
26 (0.0017)
66.29-75.27 (8.977)
50 (0.0065)
15.625-16.745 (1.121)
26 (0.0342)
26.81-31.05 (4.245)
51 (0.0001)
47.89-54.39 (6.505)
51 (0.0004)
58.82-65.26 (6.44)
47 (0.0001)
89.36-105.4 (16.05)
51 (0.0011)
11.59-13.35 (1.753)
25 (0.0176)
17.02-18.86 (1.846)
51 (0.0001)
23.53-25.51 (1.987)
24 (0.0036)
34.96-38.66 (3.703)
28 (0.0001)
46.17-54.47 (8.296)
51 (0.0001)
58.19-61.94 (3.745)
51 (0.0001)
65.23-67.12 (1.897)
28 (0.0005)
41.46-49.56 (8.100)
51 (0.0011)
36.82-41.39 (4.565)
33 (0.0346)
69.22-77.83 (8.614)
38 (0.0126)
20.68-32.60 (11.93)
51 (0.0009)
51.02-54.06 (3.033)
40 (0.0265)
85.24-85.28 (0.034)
37 (0.0498)
9.072-13.53 (4.461)
50 (0.0017)
33.32-36.43 (3.108)
38 (0.0022)
40.36-46.85 (6.481)
35 (0.007)
48.98-50.50 (1.522)
8 (0.0451)
9.24-12.36 (3.119)
32 (0.0457)
49.99-54.41 (4.412)
41 (0.0128)
25.14-29.43 (4.283)
51 (0.0001)
32.87-35.72 (2.857)
51 (0.0008)
40.02-41.82 (1.798)
51 (0.0401)
44.62-54.41 (9.792)
51 (0.0001)
63.36-64.95 (1.594)
51 (0.0051)
4.040-6.339 (2.299)
47 (0.0008)
13.97-18.06 (4.093)
37 (0.0016)
45.09-50.35 (5.261)
51 (0.0174)
65.61-66.89 (1.279)
30 (0.0324)
32.32-38.56 (6.235)
47 (0.0137)
43.79-53.04 (9.249)
34 (0.0451)
25.59-30.12 (4.529)
27 (0.0235)
52.78-57.25 (4.466)
51 (0.0255)
27.73-30.38 (2.653)
51 (0.0102)
36.33-38.47 (2.142)
37 (0.0013)
41.40-42.52 (1.115)
50 (0.0278)
Functional annotation of regions under selection
The 36,320 SNPs were representative of about 7,177 different genes corresponding to approximately one third of the total number expected in the genome. For each of these, Table S5 (additional file 2) summarizes the number of SNPs they contained and results from the differentiation analysis. Individual SNP information made it nevertheless difficult to propose a list of genes subjected to selection and this strategy might be more sensitive to possible false positives introduced by hierarchical structure among the populations under study [24]. Alternatively, a great proportion of such genes could include SNPs appearing as significantly differentiated as a result of hitchhiking with a favourable variant located nearby (see above). Hence, among the 191 (853) genes containing or close to at least one SNP with a BF > 30 (BF > 15), 103 (237) mapped within one of the 53 regions identified above. For functional and network analyses, we thus decided to mainly concentrate on the 42 genes which were located for 25 of them (represented in Table S5 in additional file 2), under the peak of each region, or for the remaining 17 less than 50 kb from the peak of each region (Table 3) (and not represented by any of the 36,320 SNPs surveyed). These 42 genes were viewed as the strongest candidates underlying the observed footprints of selection. Among these, 37 genes were eligible for network analysis, 29 being also eligible for functional analysis. The five remaining ones (CCDC46, CTXN3, KRTAP8-1, RNF180, TEKT3) were not eligible for any network or functional analyses.
Immune response genes under selection
Several candidate genes (such as CD79A, CXCR4, DLK1, RFX3, SEMA4A, TICAM1 and TRIM21) belonging to N directly participate in antigen recognition, a key process underlying the development of immune response. For instance, TRIM21, a newly identified type of IgG Fc receptors [27] participates to both the host anti-viral response through the modulation of IRF3 functions [28] and mediation of autoimmune diseases. Footprints of selection were recently reported in primates within another member of the same tripartite motif (TRIM) family of proteins which has antiretroviral properties [29, 30]. TICAM1 is a Toll like receptor (TLR) adapter and thus plays a role in innate immunity. It possesses, in particular, a high IFN type I inducing activity during viral infection [31]. DLK1, a member of the epidermal growth factor-like gene family, has been shown in mouse to be involved in B lymphocytes differentiation and function [32]. More specifically, other genes (such as CD79A, CXCR4, RFX3 and SEMA4A) are related to antigen presentation to T cells in the context of MHC. Although no MHC genes were present in the list analyzed, the corresponding region (#49 in Table 3) and a SNP (with BF = 17 and F ST = 0.005) within the bovine HLA-DMB ortholog (Table S4 in additional file 2) were found under strong balancing selection. Among the molecules participating to antigen presentation, the chemokine receptor CXCR4 is co-recruited with CCR5 in the immunological synapse, and it participates to modulation of T cells response [33]. These two molecules were shown to act as co-receptors for HIV entry in human cells [34]. A strong signature of balancing selection was reported in the 5' cis-regulatory region of CCR5 in human [35], this region displaying as well a selective sweep in chimpanzee [36]. We also recently observed a significant signal of selection on a microsatellite located less than 1 Mb from CXCR4 in West African taurine [10] and defining the boundary of a QTL underlying the trypanotolerance trait in cattle [37]. Similarly, RFX3 (43.235-43.563 Mb on BTA08) is located within another QTL identified in this latter study [37]. This member of the regulatory factor X (RFX) gene family participates with CIITA in the regulation of MHC genes [38–40]. CD79A via CD79α-CD79β heterodimer located on the surface of B cells is required for antigen presentation through MHC class II [41]. SEMA4A a semaphorin expressed on dendritic cells and B cells is involved in T cell priming and Th1 differentiation through its interaction with Tim2 expressed on T cells [42].
Although heat stress response may have imposed immunological parameter modifications [5], infectious and parasitical diseases are likely to have represented selective pressures acting on genes involved in such functions [43]. As previously mentioned, trypanosomiasis represents one of the well known examples of such selective pressure. Nevertheless, the importance of other diseases such as tick-borne diseases (anaplasmosis, babesiosis, cowdriosis) or anthrax should not be overlooked. For instance, anthrax is hyperendemic (especially in Chad, Togo and Ivory Coast) or endemic in West Africa http://www.vetmed.lsu.edu/whocc/mp_world.htm. Notice that the ANTRXR2, a receptor of the anthrax toxin was among our candidate genes (see network N, Figure 6).
Neural genes under selection
In addition to the exposure to new pathogens, the recent settlement of cattle to Sub Saharan Africa might have imposed a dramatic change in their environmental conditions [5]. Hence, cattle (in particular taurine) were exposed to warmer temperature, to a thermal amplitude decrease, to a day length change, to differences in solar radiation and also to new feeding conditions. As a result, adaptations needed modifications in global appearance (such as morphology or coat color), in body temperature regulation, in circadian clock and also in reproductive behaviour and abilities. Such selective pressures might partly explain the physiological functions related to nervous system (neurogenesis and eyesight). Alternatively, neurotropism of several parasites such as Trypanosoma(sp) have been demonstrated at least in human. Some of the candidate genes related to nervous system development and function belonged to N3 (e.g. NEUROD6 and OLFM2) or N (e.g. MAGI1, SEMA4A, HTR4 and EDNRB). NEUROD6 is a member of the neurogenic differentiation transcription factor family [44] while OLFM2 is a secreted glycoprotein belonging to the noelins family which modulate the timing of neuronal differentiation during development [45]. Interestingly, MAGI1 is a scaffolding protein present in tight junction of epithelial cells [46], some transcripts of the corresponding gene being only expressed in brain. Moreover it is located within a QTL underlying the trypanotolerance trait [37]. Besides their role in immune response (see above), semaphorins represented by SEMA4A were initially characterized for their role in the guidance of axonal migration during neuronal development. Mutations in the conserved semaphorin domain of SEMA4A are associated with two retinal degenerative diseases, retinis pigmentosa and cone-rod dystrophy [47]. Of more particular interest is HTR4 which is a serotonin receptor participating to the serotonergic system. Recently, a mutation within HTR2A, a receptor of the same family was reported associated with the chronic fatigue syndrome corresponding to a dysregulation of hypothalamic-pituitary-adrenal (HPA) axis and serotonergic system [48]. A reduced responsiveness of the HPA axis after corticotropin-releasing hormone (CRH) challenge was observed in Boran cattle infected with Trypanosoma congolense [49]. Taken together, a possible involvement of HTR4 in trypanotolerance/susceptibility in cattle might be hypothesized though we cannot exclude an implication of this serotonin receptor in more general aspects of circadian rythmicity [50].
Genes under selection involved in skin and hair properties
Although EDNRB plays an essential role in the development of enteric neurons, it is also involved in the development of epidermal melanocytes, both cell lineages being derived from the neural crest [51]. As an illustration, several EDNRB mutations are associated with an auditory pigmentary syndrome caused by the absence of melanocytes [52]. However, given the variety of cattle populations surveyed (Figure 1), the footprint of selection observed in the EDNRB region is more likely related to differences in coat color and more particularly to the spotting pattern (SOM, BAO and LAG) or the white color (KUR and ZFU). Indeed, EDNRB is also referred to as piebald or S locus in mouse [53] and a null mutation induce a white coat color in rat [51]. In the Hereford cattle breed, although not yet fully characterized, locus S seems responsible for white spotting pattern [53]. In addition to EDNRB involved in skin and hair pigmentation, other genes such as TRSP1 which belonged to N and KRTAP8-1 which was not eligible for network analysis play a role in hair development in human and mouse. Indeed, TRPS1 is a zinc finger transcription factor implicated in growth and trichosis, some of its variants being associated with trichorhinophalangeal syndromes [54, 55]. It is down-regulated in patients affected by hypertrichosis and in a mouse model of hypertrichosis [56]. KRTAP8-1, a keratin associated protein, plays a role in the formation of hair shafts [57].
The skin color and thickness, the hair size and sleekness and the number of sweat glands directly influence thermo-resistance of cattle living in the tropics [5]. Compared to European taurine breeds, zebus which have a higher density of sweat glands and a smoother and shinier hair coat were reported to better regulate body temperature and more efficiently maintain cellular function during heat stress [58]. Similarly, slick-haired Holstein cattle are more able to regulate their body temperature than wild-type [59]. Some skin and hair properties might also confer higher resistance to tick infestation [60, 61]. Interestingly, some genes involved in hair development also participate in cornification. Hence the observed footprint of selection within the cluster of keratin associated proteins (represented in the candidate genes list by KRTPA8-1, the closest to the peak) might thus also be related to horn morphology differences [62]. A striking example is the floating horns of KUR which could represent original adaptation facilitating swimming in their swamp living area [3]. Note that the identified region is also localized less than 5 Mb from the bovine polled locus [63]
West African cattle provide a valuable resource to better understand the genomic response to selective pressure arising in tropical conditions. This large-scale whole genome Bayesian scan for adaptive differentiation in populations representative of the current breed diversity allowed us to identify candidate genes involved in several key physiological functions. These results might open the way towards the identification of variants underlying these footprints of selection, in particular those involved in the resistance to tropical infectious diseases in cattle but also in other mammals such as human populations subjected to similar pressures.
Animal Material
Figure 1 provides information on the origin of the nine different West African populations sampled in our study and representatives of the three main types of cattle found in this area. Longhorn taurines were represented by N'Dama individuals from two distinct origins (ND1 and ND2). ND1 individuals (n = 14) originated from the Gaoua herd constituted by acquisitions coming from different villages in the Pays Lobi (Burkina Faso). ND2 (n = 17) samples were collected in the Samandeni herd (Burkina Faso) which was constituted in the early eighties by acquisitions from the Marahoue ranch (Ivory Coast) and originated from northwestern villages from Ivory Coast where zebu had not been introduced [64]. Shorthorn taurines were represented by three different breeds: Baoulé (BAO) samples (n = 29) originated, as ND1, from the Gaoua herd [64], Somba (SOM) individuals (n = 44) were sampled in the breed birthplace across the border near Nadoba (Togo) and Boukoumbe (Benin), and Lagune (LAG) individuals (n = 44) were sampled in the Porto Novo district (Benin) [65]. West African zebus were represented by two populations: Sudanese Fulani or ZFU (n = 43) which was sampled in the Malanville region in Benin [65] and Choah zebus or ZCH (n = 59) which was sampled in the Bol district (Chad) [64]. This latter population was initially sub-divided into two breeds (M'Bororo and Choah Zebus), however, as previously shown [10] and confirmed in our analysis with a dense marker set, no clear differentiation appeared among these. Finally, two hybrid populations were considered: Borgou or BOR (n = 45) which is a stabilized crossbred between shorthorn taurines (LAG or BAO) and was sampled in its region of origin around Parakou in Benin [65] and Kuri or KUR (n = 47) which was sampled from different islands of Lake Chad around Bol [64]. This latter breed was sometimes referred to as a particular longhorn taurine [3], however, several molecular analyses showed a high level of Zebu admixture in it [4, 10]. Moreover, unlike West African taurines, KUR is known to be trypano-susceptible. Three other breeds were considered as outgroups for the detailed genetic structure analysis: ZMA (n = 35) sampled in the Madagascar Island [66] and representing pure zebu [1], AUB (n = 20) sampled in the birthplace of the breed [67] and representing European taurine, and OUL (n = 40) sampled in the North of Morocco to represent North African taurine cattle.
Genotyping data
Individuals were genotyped on the Illumina BovineSNP50 chip assay [9] at the Centre National de Génotypage (CNG) platform (Evry, France) using standard procedures http://www.illumina.com. Among the 54,001 SNPs included in the chip, only the 51,581 mapping to a bovine autosome on the latest bovine genome assembly Btau_4.0 [68] were retained for further analysis. To limit ascertainment bias favouring SNPs from European origin, we subsequently discarded 13,786 SNPs (~25%) which were not polymorphic (MAF > 0.01) in at least one West African taurine (ND1, ND2, LAG, SOM or BAO) and one West African zebu (ZCH or ZFU). Among the remaining ones, 1,422 SNPs which were genotyped on less than 85% of the individuals from at least one of the nine West African breeds, were also eliminated. An exact test for Hardy-Weinberg Equilibrium (HWE) [69] was subsequently carried out within each breed separately. Based on the obtained p-values, q-values [70] were estimated for each SNP using the R package qvalue http://cran.r-project.org/web/packages/qvalue/index.html. Fifty three SNPs with a q-value < 0.01 in at least one breed were then discarded from further analysis. In total, 36,320 SNPs were finally considered for the study leading to an average marker density of 1 SNP every 70 kb over the genome (Table S8 in additional file 2). Moreover, as shown in Figure S6 (additional file 1) and detailed in Table S8 (additional file 2), the genome coverage was homogeneous with a median distance between consecutive SNPs equal to 47.54 kb. Few large gaps between SNPs were present since the 95th (99th) percentile of this distance was 189 kb (385 kb), the largest gap localized on BTA10 being 2 Mb long. Conversely, less than 0.5% of the distances between successive SNPs were shorter than 20 kb. Genotyping data are given in additional file 3.
Analysis of Population Structure and characterization of the extent of LD
ASD were computed for each pair of individuals using all available SNP information by a simple counting algorithm: for a given pair of individuals i and j, ASD was defined as 1-x ij where x ij represents the proportion of alleles alike in state averaged over all genotyped SNPs. A neighbor-joining tree was computed based on the resulting distance matrix using the R package APE [71]. We subsequently performed a PCA based on all available SNP information using the SMARTPCA software package [14]. As suggested by Patterson et al. [14], we performed LD correction by replacing individual SNP values with the residuals from a multivariate regression without intercept on the two preceding SNPs on the map, providing they were less than 200 kb apart. Results were further visualized using functionalities available in the R package ade4 [72]. The global F-statistics F IT , F ST and F IS were estimated respectively in the form of F, θ and f [73] using the program GENEPOP 4.0 [74]. GENEPOP 4.0 was also used to estimate diversity for each locus and population both within individuals and among individuals within a population. The within breed F IS was derived from the average of these two quantities over all the SNPs. In order to evaluate the reliability of the F IS estimates we computed the mean and standard deviation over 10,000 samples of 5,000 randomly chosen SNPs. F ST statistics between populations were estimated using both SMARTPCA [14] and GENEPOP [74] which are based on two different models of population divergence.
In order to characterize the extent of LD, we computed the r2 measure [23] between each marker pair within each breed separetely using Haploview 4.1 [75].
Bayesian Model to analyze Differentiation among SNPs
Individual genotyping data were modelled according to the reparameterized extension, recently proposed by Riebler et al. [8], of the initial Bayesian hierarchical model developed by Beaumont and Balding [7]. Briefly and in the bi-allelic SNP case, let a ij be the observed reference allele (defined arbitrarily) count in population j = 1, ..., J at locus i = 1, ..., I. The conditional distribution given the true (unobserved) allele frequency p ij in that population at that locus is assumed to be binomial with parameters n ij (twice the number of genotyped individuals in population j at locus i) and p ij : a ij | p ij ~Bin(n ij , p ij ) (1). Note that 1) implicitly assumes populations are in HWE or their respective inbreeding coefficients (F IS ) are null. Non null F IS could be taken into account in the model by considering instead that the three possible genotypes are drawn from a multinomial distribution with parameters corresponding to the number of individuals genotyped and genotype probabilities: (1-F IS j )p ij 2 + F IS j p ij ; 2(1-F IS j )p ij (1-p ij ) and (1-F IS j )(1-p ij )2 + F IS j (1-p ij ) [18]. Nevertheless, for co-dominant markers such as SNP and given the range of F IS values in our study (see Results), the binomial distribution seems to be reasonable (Figure S3 in additional file 1). The second step assumes that the true allele frequencies p ij are themselves sampled from a Beta distribution: p ij | λ ij , x i ~Beta(λ ij x i , λ ij (1-x i )) (2). This distribution relies on an infinite Wright island model involving mutation, drift and migration at its equilibrium state [76, 77]. Under this model, x i might be interpreted as the frequency of the chosen reference allele at locus i in the gene pool from which each allele frequency p ij is sampled. The scaling parameter λ ij reflects the gene flow from the gene pool to population j. Under Wright's model, this parameter is specific to each population j but remains homogeneous over loci so that any departure from that property may indicate that locus i is no longer neutral. Note that under the Beta distribution, the first two moments can be simply expressed as E(p ij ) = x i and Var(p ij ) = x i (1-x i )(1+λ ij )-1. Actually (1+λ ij )-1 can be identified as a F ST ij coefficient [17]. The next level of the model consists of specifying the distributions of λ ij (or F ST ij ) and of x i . These frequencies are nuisance parameters and uncertainty about them will be taken into account by assigning to them non informative prior such as a Beta(1,1) distribution (e.g. [78]). Following Beaumont and Balding [7], the λ ij 's are modelled via a linear model on the logistic transformation of the F ST ij . Since F ST ij /(1-F ST ij ) = 1/λ ij , we can write this model in terms of η ij = log(F ST ij /(1-F ST ij )) = -log(λ ij ) as: η ij = α i + β j + γ ij where α i is a locus effect, β j a population effect and γ ij an error term corresponding to a departure of the logit of F ST ij from the additive decomposition. For theoretical and computational reasons, it is more efficient to consider the previous decomposition under a hierarchical Bayesian structure than under its basic form, and implement it as follows: η ij | α i , β j , σ γ 2~ iid N(α i + β j , σ γ 2) with β j | μ β , σ β 2~ iid N(μ β , σ β 2) and α i | σ α 2~ iid N(0, σ α 2). Introduction of additional levels by defining priors on the variance components σ α 2, σ β 2 and σ γ 2 is straightforward but adds marginal gain in the estimates and requires high supplementary computational costs (Gautier and Foulley, unpublished data). We thus gave for these components the known values proposed initially by Beaumont and Balding (2004): σ α 2 = 1, σ β 2 = 3.24 and σ γ 2 = 0.25. The prior mean μ for the β j was also taken as -2.0. Recently, Riebler et al. [8] introduced in the above logistic model an auxiliary indicator variable δ i attached to each locus specifying whether it can be regarded as selected (δ i = 1) or neutral (δ i = 0). They demonstrated the efficiency of this approach for improving the power of the statistical procedure, particularly for the detection of loci under balancing selection. Under this reparameterized model, the previous parameters α i are written as: α i = δ i α i * with α i * | σ α *2~ iid N(0, σ α *2). The model further assumes a Bernoulli distribution for the indicator δ i variable with parameter P: δ i | P ~Bin(1, P). P is itself assumed to be Beta distributed to take into account uncertainty on this crucial parameter. Here we took P ~Beta(0.2,1.8) [8], thus assuming that a priori 10% of the loci are on average under selection, but within a very large range of possible values as the 95% credibility interval goes from almost 0 to 65%. Note that the value σ α *2 can be derived from σ α 2 since by construction σ α 2 = E(P) σ α *2 (hence σ α *2 = 10).
The posterior distributions of the different parameters of interest were estimated via MCMC procedures as previously described [8] from 10,000 post burn-in samples (with a burn-in period of 3,000 iterations) and a thinning interval of k = 25. Convergence was checked using standard criterion. Parameter estimates for the F ST ij were taken as the median of the posterior distribution.
Decision rule to identify non neutral loci
where π(P) is the density of the distribution of P defined above. To make interpretation of the BF i easier, we expressed them in deciban units (dB) ie dB i = 10log10(BF i ) so that 10 dB corresponds to an odds ratio of 10, 20 dB to an odds ratio of 100, 30 dB to an odds ratio of 1,000 and so forth. We then applied the Jeffreys' rule [79] which quantifies the strength of evidence (here to consider a SNP as being under selection) based on BF using the following scale: "strong evidence" when 10<dB i <15, "very strong evidence" when 15<dB i <20 and "decisive evidence" when dB i >20. Notice that the BF captures the change in the odds in favor of δ i = 1 as we move from the prior to the posterior. In that way, it makes this criterion independent of the prior distribution of δ i and thus guarantees some robustness.
Simulated data
Following Foll and Gaggiotti [20], four data sets each comprising 50,000 (unlinked) SNPs with respectively 9300 (P = 0.186), 5000 (P = 0.1), 500 (P = 0.01) and 50 (P = 0.001) under selection (α i ≠ 0) were simulated under the inference model. To mimic characteristics of our samples, we considered 9 populations with parameters β j equal to the ones estimated on the real data (mean of the corresponding posterior distribution). For each pair of simulated locus/population, we subsequently simulated the η ij parameter by adding to the corresponding β j , a value of γ ij sampled from a Gaussian distribution (γ ij ~N(0,0.25)) and a value for α i equal to 0 if the locus was neutral or sampled from a Gaussian distribution (α i ~N(0,10)) if the locus was under selection. The frequency x i in the ancestral population (or migrant gene pool) was sampled from a Beta distribution with parameters equal to 0.7. This is equivalent to assume the ancestral population was at equilibrium with 4Nμ = 0.7 where N is the effective population size and μ the mutation rate [80]. Based on these simulated values for x i and η ij we subsequently derived the p ij parameter of the binomial distribution from which the simulated observed a ij was sampled (taking n ij = 80 for all i and j). Only SNPs displaying MAF>0.01 in at least one simulated West-African taurine and one West-African zebu (identified according to the simulated β j ) were conserved which led to the exclusion of few SNPs (displaying α i too high in general). To a first approximation, this also allowed the mimicking of the ascertainment bias introduced by the SNP selection criterion. The posterior distributions of the different parameters of interest were then estimated as described above from 1,000 post burn-in samples (with a burn-in period of 3,000 iterations) and a thinning interval of k = 25. BF i for each simulated loci were then computed (see above) and used to compute the (true) FDR and FNR for a given threshold. To investigate relationships between the coefficient of selection and the magnitude of the locus effect α i we performed simulations under a simplified version (without mutation) of the Wright-Fisher island model with selection proposed by Beaumont and Balding [7]. Briefly, 10 random mating populations with a constant size of 1,000 individuals and deriving from the same ancestral population were reproduced for 500 generations assuming a global F ST of 0.15. In total, 5,000 neutral SNPs and 500 SNPs for each investigated coefficient and mode of selection were simulated. Initial allele frequencies were drawn from a Beta distribution with both parameters equal to 0.7. At the end of the process, SNPs displaying a MAF <0.01 in at least 2 populations were discarded. Locus specific F ST were then computed following Weir and Cockerham [73]. Following Foll and Gaggiotti [20], for each type of selection considered, the corresponding average F ST was compared to the one obtained under neutrality (α i = 0) to estimate the average SNP effect.
Annotation of the SNPs
Because the annotation of the bovine genome is still sparse, the gene content information was derived from the TransMap cross-species alignments available in the UCSC Genome Browser http://genome.ucsc.edu/. For closer evolutionary distances, the alignments are created using syntenically filtered BLASTZ alignment chains, resulting in a prediction of the orthologous genes in cow. In total, 46,598 different RefSeq identifiers were anchored to the Btau_4.0 bovine genome assembly http://genome.ucsc.edu/. Considering that most consecutive SNPs on the map were separated by more than 20 kb and the relatively high level of LD at shorter distance (see Results and [11]) a SNP was considered as representative of a gene if it was localized within the boundary positions of the gene that extended by 15 kb upstream and downstream. According to this criterion, 15,360 out of the 36,320 SNPs were representative of 17,190 different TransMap RefSeq identifiers. Subsequent annotation and analyses were carried out with the Ingenuity Pathway Analysis (IPA) software v7.0 (Ingenuity Systems Inc., USA, http://www.ingenuity.com/). Among the 17,190 different TransMap RefSeq identifiers, 17,151 identifiers (99.7%) were represented in the Ingenuity Pathway Knowledge Base (IPKB) and corresponded to 7,177 different genes further considered as the reference set. Finally, 15,336 SNPs were located within a gene, leading to an average of 2.21 SNPs per annotated gene. Notice that 421 of these SNPs were representative of more than one gene.
Functional and Network Analyses
Functional and Networks analyses were carried out using an approach similar to the one described in [81]. Based on the information contained in the IPKB, IPA allows performing both functional and network analyses. The functional analysis aims at identifying the most significant biological functions represented in the candidate gene list from the reference set. Note that among the genes under selection, only those associated with at least one functional annotation in IPKB were eligible for functional analysis. The most significant functions are then obtained by comparing functions associated with eligible genes against functions associated with all the genes in the reference set using the right-tailed Fisher's exact test. The network analysis aims at searching for interactions (known from the literature) between candidate genes and all other IPKB molecules (genes, gene products or small molecules) and result in the definitions of networks which contains at most 35 molecules (including candidate genes). For each network, a score S is computed based on a right-tailed Fisher exact test for the overrepresentation of candidate genes (S = -log(p-value)). In our study, networks were considered as significant when S>3 and their associated top functions and diseases were further reported.
Identification of regions under selection
In order to identify regions under selection (with an unexpectedly high proportions of SNPs subjected to selection), we followed the locally adaptive procedure which allows to account for variations in distance between the different tested positions [81, 82]. Individual SNP BF values were first smoothed over each chromosome with a local variable bandwidth kernel estimator [83]. We further performed 250,000 permutations to estimate local p-values which were corrected for multiple testing by computing q-values (see above).
List of Abbreviations
- Breed Codes :
AUB: Aubrac
N'Dama (first population)
N'Dama (second population)
Oulmès Zaer
Choah zebu
White Fulani Zebu
Zebu from Madagascar.
: Allele Sharing Distance
Bayes Factor
Bos TAurus chromosome
- dB:
deciban unit
Hardy-Weinberg Equilibrium
Ingenuity Pathway Analysis
Ingenuity Pathway Knowledge database
False Discovery Rate
False Negative Rate
Linkage Disequilibrium
Minor Allele Frequency
Major Histocompatibility Complex
Principal Component Analysis
Single Nucleotide Polymorphism.
The authors wish to thank the farmers, the persons in charge of breeding programmes and the CIRDES (Burkina-Faso) team for their precious help in providing the samples from Burkina-Faso, Togo and Benin and Pr Lahoussine Ouragh from the Rabat Institute of Morocco for providing samples from Oulmès-Zaer breed. They also wish to thank anonymous referees for helpful corrections. The genotyping was supported by the French Ministry of Education and Research and the National Institute of Agronomic Research (INRA AIP "Bio-Ressources").
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This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


text:
Glen Williams wrote:
> Ran ,
> I applaud your efforts and those of other hybrizers to get more of a
> return for the time and effort ( AND RECORD KEEPING) you put into your
> work. The limited TC run may make the initial buyer a little happier and
> limit conspicious consumption for a little while , but surely you still
> face the problem of someone buying an early TC plant from you and in turn
> creating a thousand plants from that one and flooding the market.Other than
> a degree of moral pressure what is there to stop the "in-for-the-buck" TC
> er for capitalizing on your limited edition?
> In addition do I understand that you will not be offering originators
> stock? Just TC. OS always seems to be a lure for some of us.
> Glen
> Glen Williams Zone 5a Springield , Vermont.
> "...and I am unanimous in that."
> ---------------------------------------------------------------------
I will continue to offer OS material. I am thinkubg however, that the
Tc labs may not be so ready to "pick up" some of the Hostas that remain
unreistered as the market for these may be to limited with the new AHS
rules. This whole thing may not work just as the plan was designed. I
will wait and see!
To sign-off this list, send email to email@example.com with the
message text UNSUBSCRIBE HOSTA-OPEN

subset: commoncrawl


text:
Category:Karin Schäfer Figuren Theater
From Wikimedia Commons, the free media repository
English: Karin Schäfer, born in Mödling, Austria, September 21, 1963 is a performance artist and the head of the Karin Schäfer Figuren Theater - Visual Theatre Productions company
This category has only the following subcategory.
- ► Karin Schäfer 'Wind and Beyond' (2 F)
Media in category "Karin Schäfer Figuren Theater"
The following 27 files are in this category, out of 27 total.
- Karin-schaefer-workshop.jpg 974 KB


text:
Not sure. Are all plants prominently labeled? I think that would be better
than a handout. If you wanted to give them a list, perhaps you could do a
simple graph type thing like my local phone book does for a city map.
Section your garden off in say 3 sections by 3 sections for 9 sectors and
say that the Ilex opaca is in B-3. It'll get them close enough to the
location to find the label. Mind you, I don't mean you'd physically label
the sections in the garden itself. B-2 would obviously be the center area
as opposed to C-3 being the southeast corner. That labeling would just be
on the paper handout.
neIN, Z5
----- Original Message -----
From: "Donna" <email@example.com>
To: <email@example.com>
Sent: Sunday, February 20, 2005 12:07 PM
Subject: RE: [CHAT] garden documentation
> interesting... I really need to get this done soon. As
> some of you know, my yard will be on a pre-tour for DH
> international group... although they will be looking
> at the trains (ha! he better get working on them!), my
> thought was to have some kind of handout with the
> plants listed by location. Probably never happen in
> time, but ......
> Hey Kitty.... what is the easiest way to do this?
> Donna
> --- email@example.com wrote:
> > Donna,
> >
> > What information I include for my garden rooms is
> > fairly basic: cultivar
> > name and common name. I don't need to add location
> > to these documents as
> > they are organized by location.
> ---------------------------------------------------------------------
> To sign-off this list, send email to firstname.lastname@example.com with the
> message text UNSUBSCRIBE GARDENCHAT
To sign-off this list, send email to email@example.com with the
message text UNSUBSCRIBE GARDENCHAT


text:
Security firm Symantec tells the Senate Congressional Committee that Stuxnet was a wake-up call for cybersecurity experts.
The malware was known for its complexity. Stuxnet is still a bit of a mystery with speculation that an Iranian uranium enrichment facility was a target.
CNET reports that the worm was the first known threat to spy on and reprogram industrial control systems and grant hackers control of critical infrastructure.
This story is part of Federal News Radio's daily Cybersecurity Update brought to you by Tripwire. For more cybersecurity news, click here.
Connolly calls for 3.2 percent pay raise for feds in 2018
Connolly calls for 3.2 percent pay raise for feds in 2018
Photos of the amazing, moving, important and amusing things happening in the federal community.
A daily update of important moments in the history of the U.S. government.


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REAL MADRID believe that a bid of £30 million would prompt Manchester United to part company with David Beckham, but are prepared to offer an unidentified player as a makeweight in what would be nonetheless a record transfer deal for a British footballer. No formal contact has been made between the two clubs, but Real are increasingly confident that an agreement will be reached with Beckham and his club.
The figure of £38 million has been widely quoted as United's valuation of Beckham, but Real believe that the deflated transfer market, added to the England captain's reluctance to discuss an


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02.27.2017 Swirling Dust in Gale Crater, Mars, Sol 1613
02.27.2017 Dust Devil Passes Near Martian Sand Dune
02.27.2017 Sand Moving Under Curiosity, One Day to Next
02.08.2017 Mars Reconnaissance Orbiter Observes Changes
01.26.2017 Mono Lake
01.25.2017 'Wing' Dike of Hardened Lava in New Mexico
01.25.2017 Blade-Like Martian Walls Outline Polygons
01.23.2017 Spirit And Opportunity By The Numbers
01.10.2017 Mars 2020 Rover - Artist's Concept
01.06.2017 Earth and Its Moon, as Seen From Mars
12.13.2016 Now and Long Ago at Gale Crater, Mars
12.13.2016 Where's Boron? Mars Rover Detects It
11.15.2016 Schiaparelli Impact Site on Mars, Stereo
11.03.2016 Schiaparelli Impact Site on Mars, in Color
10.17.2016 MAVEN Captures Rapid Cloud Formation
10.17.2016 Mars' Nightside Atmosphere
10.17.2016 Ultraviolet Image Near Mars' South Pole
10.17.2016 Ultraviolet Mars Reveals Cloud Formation
10.05.2016 Dust Haze Hiding the Martian Surface in 2001
10.04.2016 Test of Lander Vision System for Mars 2020
10.03.2016 A Sharpened Ultraviolet View of Mars
10.03.2016 Curiosity Self-Portrait at 'Murray Buttes'
10.03.2016 Butte 'M9a' in 'Murray Buttes' on Mars
09.19.2016 Ribbon Cutting
09.09.2016 Farewell to Murray Buttes (Image 5)
09.09.2016 Farewell to Murray Buttes (Image 4)
09.09.2016 Farewell to Murray Buttes (Image 3)
09.09.2016 Farewell to Murray Buttes (Image 2)
09.09.2016 Farewell to Murray Buttes (Image 1)
08.26.2016 Out-of-this-World Records
08.04.2016 Mars Rover Is New Social Media Game
08.04.2016 Mars Rover Social Media Game
08.02.2016 Artist Concept for RIMFAX
07.20.2016 Viking 40 Year Anniversary Artwork: Medal
Weighing Molecules on MarsThe plot on the left shows new results from the Sample Analysis at Mars, or SAM, instrument on NASA's Curiosity rover. The instrument measured levels of different gas isotopes in the atmosphere. Isotopes are variations of atoms weighing different amounts. As seen on the plot, SAM detected about 2,000 times as much argon-40 as argon-36, which weighs less. This result is the most precise measurement yet of argon isotope ratios on Mars, and confirms the connection between Mars and Martian meteorites found on Earth, an example of which is shown at the right. The dark blobs in the meteorite are areas where atmospheric gases were trapped when the meteorite was ejected from Mars, and they include argon with the same ratio of argon-40 to argon-36 as SAM has measured in Gale Crater.
The data at left also show peaks for carbon dioxide containing different isotopes of carbon and oxygen. The carbon dioxide at mass 44 contains the most abundant isotopes of both carbon and oxygen, so it has the largest signal, with its peak being higher than the top of the chart shown here.
Image Credit: NASA/JPL-Caltech/JSC
All Images


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Home > All Cities > Marietta, GA Real Estate Listings > Dorset Subdivision
Properties for sale in the Dorset Subdivision, Marietta, GA
Properties 1 to 1 of 1
3145 Cherbourg Ct, Marietta, GAList Price: $435,000 | Bed:4 | Bath:3.10
Beautifully maintained home in Dorset, a swim/tennis community located in the Pope H.S. district. Located at the end of a cul-de-sac, this property has an updated master bathroom with granite, ample counter-space and storage. New windows and skylights, 2 year old architectural roof, garage doors and openers, keyless entry. Ready for you to enjoy! Cul De Sac, Sloping, Level Lot 50x170x25x199x151 Bookcases, Cable TV Connections, Ceilings - Trey, Ceilings - Vaulted, Double Vanity, Foyer - 2 Story, Foyer - Entrance, Garden Tub, Hardwood Floors, Recently Renovated, Separate Shower, Skylight, Tile Bath, Tile Floors, Walk-in Closet, Wall-to-wall Carpet Neighborhood Association, Playground, Pool, Swim Team, Tennis Courts, Tennis Team, Underground Utilities, Walk To Schools Deck/Patio, Garden Area, Sprinkler System, Workshop
Listing Office: KELLER WILLIAMS REALTY CONSUL | MLS ID#: 07623076


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Fujitsu introduces ARROWS Z ISW11F au smartphone
Company [Thursday 15 December 2011]
Fujitsu announced that the new ARROWS Z ISW11F smartphone from Fujitsu Toshiba Mobile Communications will be available in Japan beginning December 17, 2011, from KDDI Corporation and Okinawa Cellular Telephone Company. To deliver unprecedented speed of operations, ARROWS Z ISW11F combines three high-speed enhancements - high-speed WiMAX communications, a high-performance 1.2GHz dual-core CPU, and a quick-response touchscreen.
Category: Product launch | Posted: Dec 15, 10:23 | More info
DAF 2014 Smart City Forum
Computex Taipei 2016
Taiwan Photonics Festival 2015
Computex Taipei 2015
Simbe Robotics's Tally solution
Asustek G11 gaming desktop
Asustek ZenFone 3 Zoom smartphone
Asustek ZenFone AR smartphone
Gorilla Glass concept car
Intel Compute Card
Asustek Zenbo home service robot
VIA Artigo A830 IoT kiosk solution
Samsung 14nm chips for wearables


text:
10 Groups to Join on LinkedIn if You're in Marketing
With over 450 million users in 200 countries, LinkedIn is the world's largest professional network. From growing your business connections, learning best practices, and staying up to date on trends, LinkedIn is a one-stop-shop for marketers looking to advance their skills and career. Without the noise of other platforms, LinkedIn strives to maintain a (mostly) professional standard.
8 (of the) Best Web Analytics Guides of 2010
Which tools and plugins are really useful for extending the capabilities of Google Analytics (GA)? What common analytics mistakes should you avoid? Discover the answers to these questions and others here in eight of the best guides to web analytics of last year. Simple Google Analytics Tips You Should Be Using by The Daily SEO Blog. Post on Google Buzz. Digg this!
Big Data is Buzzing, But Small Data Packs a Punch [Q&A]
Sales Intelligence View
But as both sales and marketing organizations recognize a clear need to understand their prospects at a deeper, more analytical level, these teams are starting to employ a holistic data strategy. He was founder and CEO of one of the first SaaS-based web-analytics companies, Keylime Software (Yahoo!), and he led products and strategy at Webtrends. But it is now. It's wider at the top.
Best SEO Guides, Tips and Resources of 2010
Frequent best-of contributor Stoney deGeyter elaborates on the "key components to a well-rounded optimization campaign, including keyword research, website architecture, copywriting, on-page optimization, analytics and more. How to optimize page Titles and improve click through rates by Web SEO Analytics. 40 (of the) Top SEO Guides, Tips and Resources of 2010 (So Far).
55 (of the) Best Social Media Tips, Tactics and Tools of 2010
Dan Neely discusses which social networking sites get the most attention from marketers (Facebook, Twitter, LinkedIn, no surprises there), how marketers can best use social media for branding and business development, and concerns about the use of social media in brand planning. Which Profile Aspects Should Be Emphasized on Twitter, Facebook and LinkedIn? by Green Buzz Agency.
18 Ways to Promote Your White Paper for Lead Generation
This is an ideal method to generate back links and create buzz about your paper. Write a short paragraph on what your paper is about (think about the target audience) and get it out on LinkedIn groups, Twitter, Facebook, and Google+. With LinkedIn, promote your white paper in a form of discussion. . Well, your firm has written and produced a white paper. Generate new leads.
87 More Vital Social Media Marketing Facts and Stats for 2012
and LinkedIn is the "oldest" network, with 79% of users age 35 and older. Slashdot is almost 90% guys); LinkedIn is close to a 50-50 split; Twitter and Facebook are both about 40% male / 60% female; and women account for almost 90% of Pinterest users. Digital Buzz Blog ). Digital Buzz Blog ). Digital Buzz Blog ). Digital Buzz Blog ). Pingdom ).
17 Ways to Promote Your White Paper for Lead Generation
This is an ideal method for generating backlinks and creating buzz about your paper. That's Post it out on LinkedIn groups, Facebook, and Google+. On LinkedIn, promote your white paper in the form of a discussion, for example: "How has technology improved the performance of your sales staff? Many association websites have resource sections. Find That was a lot of work.
Why Modern Marketers Fail at Big Data
Sales Prospecting Perspectives
Without the right analytics and a targeted campaign, all they've got is a huge number of eyes. All of these examples demonstrate valuable examples of industry-leading brands with huge budgets and accessible resources that do not understand how to derive real value from their marketing campaigns. Imagine the sorts of campaigns their marketing teams could have run with these captive audiences had they understood their best customers better instead of simply sitting back and counting the voices buzzing about their brands. Find her on Google+ or LinkedIn Duck Dynasty.
Our Top 10 Blogs of 2016 at a Glance
Modern B2B Marketing
Ways to Boost B2B Sales Through LinkedIn Social Selling. Ways to Boost B2B Sales Through LinkedIn Social Selling. With more than 400 million members in over 200 countries and territories, LinkedIn is an effective way to reach an audience with a business mindset. LinkedIn is also a great channel for sales teams to drive revenue with. His recommended tactics?
How to Create Irresistibly Shareable Content
KoMarketing Associates
By mentioning key thought leaders and publications in the industry, you can generate social buzz and build relationships for potential guest contributions and other link building tactics. would highly recommend searching keywords on popular forums like Quora , exploring keywords and phrases in Twitter search and seeing what people are asking in industry related LinkedIn Groups. Enjoy!
An Eight-Step Process for Creating Front-Page B2B Industry News
KoMarketing Associates
While every client is different, we have a basic framework in place for creating news articles from start to finish, which ultimately results in getting our news buzzing within the appropriate industries. Aggregating Resources. When setting up a Feedly content stream, we either browse by category or enter some keywords to identify resources that could be beneficial for news creation.
33 (of the) Best Social Media Guides, Tips and Resoures of 2012 So Far
David Rabjohns says that the top five questions Fortune 500 companies are asking about social media are: • Where are people talking about my brand? • How should I change my messaging? • How much buzz do I have vs. competition/trend? • How do they feel about us vs. the competition? 103 Resources For Becoming a Pinterest Expert by KISSmetrics. " Don't be one of those brands.
The Un-Tapped Value of Employee Social Capital
grow - Practical Marketing Solutions
Social ROI has become a hot buzz word in the industry and corporations are finally figuring out how to apply social analytics and really derive actionable insights from social media conversations. By Konstanze Alex Brown, {grow} Community Member. Why should any business owner like social media? Why should you care? What company does not want that? Let's dig in a bit. Not so much.
B2B Lead Generation Blog: Google Makes Web Analytics Free but will it help your lead generation?
B2B Lead Generation Blog
B2B Lead Generation Blog About Free Resources Get FREE Updates Instantly Get A FREE Chapter from my book. « Winning the Complex Sales Cycle with Thought leading Content | Main | Embattled CMOs share strategies to improve marketing influence » Google Makes Web Analytics Free but will it help your lead generation?
9 Reasons "Home Alone" Is Absolutely Clutch for Content Marketers
KoMarketing Associates
Do your research in Analytics, Search Console, etc. Content Marketing Takeaway: Strive to be as resourceful as Kevin was! If you weren't already team Kevin, the moment he goes to visit Santa and asks for "nothing but Peter, Kate, Buzz, Meghan, Lynny, and Jeff" could sway even the biggest Grinch. Check out the types of questions readers are asking in LinkedIn Groups and forums.
10 Questions to ask in a B2B Content Marketing Questionnaire
KoMarketing Associates
Continually monitoring these sites throughout the process can lead to new content ideas or simply keep the content team informed of what's truly buzzing in the industry at a certain point in time. 6. What are some of the leading social media resources in the industry? Can you please provide a succinct overview of your business? What are your organization's primary industry targets? .
73 Experts Reveal B2B Marketing Trends to Leverage in 2017
Dedicate plenty of resources to content promotion. so you should definitely use research tools (including Google Analytics on your site, Facebook Audience Insights, and the Facebook Ad dashboard of who is responding to your ads and where) to find out who your customer is and what devices they use. Maximise your presence on Twitter and LinkedIn to harvest relationships. Wes Yee.
Best Social Media and Digital Marketing Research and Statistics of 2011, Part 2
Twitter, Facebook And LinkedIn: Age, Ethnicity And Gender Of The Major Social Networks [STUDY] by All Twitter. Shea Bennett examines research finding that, no surprise, the user base on LinkedIn skews older than on other social networks: more than half of all members are aged 36-65, with just 6% younger than 22. Infographic: Social Media ROI Statistics by Digital Buzz. hours.
5 Proven Strategies for B2B Social Media Marketing
delicious b2bmarketing
Top Topics Twitter YouTube Facebook iPhone Google Video Google Buzz Social media Android Mobile 2.0 Increased spending on online marketing is driving companies to try new and innovative means of getting the word out about their products and services — one area that's getting a lot of buzz with B2B marketers is social media. Create a Specialized Community Reaching the correct audience is one of the most important tasks in marketing — otherwise, you're just wasting resources on the wrong people. Be a resource. 4. Learn more. Thanks, Erica.
10 of the Most Successful B2B Blog Post Themes
KoMarketing Associates
After mining through our clients' Google Analytics data, I've put together a list (in no particular order) of ten blog post "themes" that have driven an abundance of traffic to their B2B websites. Compiling lists of "best in class" social profiles provides readers with a valuable resource they can visit time and time again and could also open communication channels with said organizations.
5 Proven Strategies for B2B Social Media Marketing
delicious b2bmarketing
Top Topics Twitter YouTube Facebook iPhone Google Video Google Buzz Social media Mobile 2.0 Increased spending on online marketing is driving companies to try new and innovative means of getting the word out about their products and services — one area that's getting a lot of buzz with B2B marketers is social media. Create a Specialized Community Reaching the correct audience is one of the most important tasks in marketing — otherwise, you're just wasting resources on the wrong people. Be a resource. 4. Learn more. Don't go it alone.
30 Terrible Pieces of Social Media Advice You Should Ignore
Especially if you have limited time and resources, don't spread yourself too thin by trying to maintain an active presence on every single social media site. And as new social networks pop up (as they do all the time), feel free to experiment with them , but be ready to let them go if they don't work for you, and let your analytics be your guide. or LinkedIn. Have no fear!
The Ultimate Glossary: 120 Social Media Marketing Terms Explained
Thus, while we're all learning about social media and inbound marketing as well as teaching others about it, having a resource that quickly and clearly explains all of the basic terms and applications is critically important. Its analytics service can show you which pages are trending, where people are interacting with your brand, and what they're saying about your content on Twitter.
A Small Biz Guide to Digital Marketing Acronyms
Vertical Response
GA: Google Analytics i s a website and analytics reporting service created by Google that gives you various statistics in regards to website traffic. There's a wide variety of social media platforms like Facebook, Twitter, LinkedIn, YouTube, Pinterest, Google+ or writing your own blog. Sign up for the free VR Buzz. © 2014, VR Marketing Blog. " Say what?
33 (of the) Best Marketing Strategy Guides and Insights of 2010
As we shift resources from traditional outbound marketing to inbound attraction marketing, how should we (re)organize to support that? Bob Leonard details 14 key factors for developing an effective b2b social media plan, among them: include input from sales, develop target personas, have a realistic content development plan, and build in analytics. Post on Google Buzz.
90 Tips to Make Your Blog Rock
delicious b2bmarketing
Jeffbullas's Blog Internet Marketing Home Bio Speaking Grade your Site Marketing Resources Twitter Jeff's YouTube Contact Congratulations: You have successfully subscribed! Promote and distribute your posts on to other social media platforms such as Facebook, Twitter and LinkedIn 29. It will be a constant resource for clients as well as adding to your websites SEO.
The ABCs of Content Marketing: A Glossary of Terms
Analytics. Making use of analytics is the only way you''ll have insight into which content is working -- or falling flat. Ideally, you''ll make use of a closed-loop analytics solution so you''re able to see what happens to all that site traffic. Spend some time carefully, and analytically, picking keywords (both short and long-tail) to optimize your content and website pages.
10 Ways to Drive Instant Results With Inbound Marketing
Share educational content with relevant LinkedIn Groups. You must supplement any paid ad spend with organic content, of course, but promoting an educational resource -- like a free ebook, for example -- using Facebook ads is a great way to get your social media lead generation off the ground, and generating results fast. Add visual calls-to-action to your site.
Top 15 Recommended Social Media Books of 2010
WindMill Networking
And what a year 2010 has been: The rise of Facebook to become a dominating (and sometimes threatening) force globally, Twitter becoming a mainstream realtime communication platform embraced by older generations, and even LinkedIn adding various functionalities throughout the year to continue to remain relevant as the default social network for professionals. Socialnomics by Erik Qualman.
Blog Marketing Tactics - 10 Tested, Proven Ideas
delicious b2bmarketing
Use other social channels for distribution, but focus on your site I don't do too much to nurture followings in external networks, I simply feed content into Twitter, Facebook, LinkedIn, etc. The Future Buzz Facebook page is being used primarily as just another way for readers to get content 8. Yes, blogs are best consumed through feeds. Exactly. 6. What did I learn?
Twitter for B2B Marketing
MI6 Marketing Agency
It virtually takes little budget (unless you want some of the fancier analytic Twitter apps. Tweetreports and Company Buzz(within LinkedIn) have both helped me in addition to the common fav's. Twitter works great for "keeping in touch and getting new engagements better than Linkedin. Great sounding board/resource. Advice from Marketing Peers. Every Thursday night Jeremy Victor and a group of B2B marketers hold a B2B Twitter Chat session. These sessions are a tremendous source of ideas, insights and data. Mining For Gold in the Chatstream. People.
The 39 Social Media Tools I'll Use Today
delicious b2bmarketing
find it to be especially valuable and hassle-free for updating Twitter, Facebook, and LInkedin simultaneously. I do this only occasionally, however, as my friends/followers on each site don't have much overlap. Tracking Google Analytics No surprise, this is my primary statistical source for Convince & Convert. Twitter Tweetdeck I use Tweetdeck for serious twitter sessions.
7 Reasons Agile Marketers Are Better at Their Jobs Than You
The concept of "agile" anything is kind of used as a business babble buzz word. Agile marketers are the ones hopping into LinkedIn Answers to answer questions, tweeting breaking news content before their competitors, sharing their funny spin on the latest meme on their Facebook Page , and carrying on conversations in real time with their social networks. is it?) Think about it.
The Ultimate Glossary: 101 Social Media Marketing Terms Explained
As we all learn about social media and inbound marketing and we teach others about it, having a resource that quickly and clearly explains all of the basic terms and applications is critically important. BackType - BackType is a social media analytics company that helps companies measure their social engagement. LinkedIn - LinkedIn is a business-oriented social networking site.
50 (of the) Best Twitter Guides, Stats, Tips and Tools of 2010 (So Far)
For those unsure of what hashtags are or unaware of their many uses on Twitter, Robin Broitman offers an extensive list of hashtag resources and guides. Twitter stats and analytics tools – top 10 by Social Media Today. Twitter ROI: Show Your Clients the Effectiveness of Twitter Campaigns by Web Analytics World. Twitter Apps Worth Another Look by Web Analytics World.
How to Structure a Kick-Ass Marketing Team for Any Company
We're going to focus on those companies who are big enough to warrant a dedicated marketing resource. Analytics: Your marketer doesn't need to be an Excel junkie, but they should have a working knowledge of the marketing funnel, know what questions to ask and be comfortable doing some basic tracking and ratios using tools like Google Analytics, Excel or even HubSpot. GREAT!


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Breach Gallery
In February 2001, renowned FBI operative Robert Hanssen was found guilty of treason against America. Over a period of more than two decades, Hanssen systematically and deliberately sold his country's key intelligence to the former Soviet Union. Academy Award® winner Chris Cooper stars as Hanssen, one of the most notorious spies in the history of America. Ryan Phillippe plays Eric O'Neill, the young agent-in-training handpicked by the FBI to help draw Hanssen from his cover. Check out photos from the film…
Roll your mouse over the left and right-hand sides of the image and then click to view more pictures.
Read our review of Breach or our interviews with Chris Cooper and Ryan Phillippe and director, Billy Ray
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- The Magnificent Seven: Cowboys descend on West End - Photo Gallery


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What do you want to watch movie-wise this weekend without leaving the comfort of your warm and snuggly home? Here's a short list of movies On Demand and others you can watch free online. ON DEMAND FROM COMCAST: Watch to Squeal at Dreamy Young Men: The Twilight Saga: New Moon R-Pattz and K-Stew and T-Laut […]
Look Out, World: It's a 'Showgirls' Sequel
By: Andy Hunsaker on October 6, 2009 in The Movies
You may feel a chill in the air right now. A sense of foreboding that heralds an ominous dread for what's to come. That would be the fact that there is actual work being done to make a sequel to 1995's obnoxious and aggressively stupid career-killing disaster Showgirls. According to a German website, writer/director Marc […]
Top 4 Future Celebrity Memoirs
Top 4 Future Celebrity Memoirs
By: Ann Murray-Yavar on June 11, 2008 in General
Update: Vanity Fair Will Not Retract Gershon/Clinton Story
By: Ann Murray-Yavar on June 10, 2008 in General
News: Gina Gershon Fighting Allegations She Had Affair With Former Prez
By: Ann Murray-Yavar on June 6, 2008 in General


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Launch a youtube similar theme on my website + minor modifications
This project received 7 bids from talented freelancers with an average bid price of $272 USD.Get free quotes for a project like this
Project BudgetN/A
Total Bids7
Project Description
Upload a script unto my website
The name of the script id "Cumulus Clips" (Youtube similar) and modify the similar with some of the features on my current auction script "Sitemile.com".
The goal is to have users upload videos and have the ability to receive funding from other users based on the quality of work showcased.
As soon as the right programmer is found the work will begin.
2000 videos are in the pipeline to be uploaded. Some of the videos are over 4GB in size.
Programming & Photoshop expertise required.
Here is additional details about the youtube similar:
Download the latest stable release of the CumulusClips video sharing cms. CumulusClips is recommended for creating new video sharing websites as well as adding video sections on existing websites. Make sure your hosting environment meets the system requirements before installing the software. The installation process takes no more than 3 minutes! See this guide for more details on installing CumulusClips.
Version Downloads Date
1.3.2 zip (2.4 MB) | [url removed, login to view] (2.2 MB) 2013-Feb-23
Moving Forward
Getting support for your video sharing website is easy. We've got a community of developer experts and site administrators ready help you with your questions. Visit our community forums to get help. If you're unsure about hosting for your video website we have organized a list of supported web hosts whom we have worked closely to ensure full support for CumulusClips.
Skills Required
Looking to make some money?
- Set your budget and the timeframe
- Outline your proposal
- Get paid for your work
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Is it unbearably hot there? And are you craving for a yummy dessert, but thinking about pounds you might gain? Here is a super light and yummy yogurt parfait! What is more, the sweet flavor comes from black mulberries, no additional sugar, which is much healthier. This dessert is perfect for people with diabetes too!
Black Mulberry Jam
Whenever I see black mulberry at the market, I remember the mythological story in the book Metamorpheses written by Ovid, a Roman poet. I read this story at my university years and was so mch impressed by it. The story takes place in Babylon, where mulberry finds the suitable climate.


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2 Corinthians 12:9 - And he said unto me, My grace is sufficient for thee: for my strength is made perfect in weakness. Most gladly therefore will I rather glory in my infirmities, that the power of Christ may rest upon me.
Psalms 34:17-20 - [The righteous] cry, and the LORD heareth, and delivereth them out of all their troubles. (Read More...)
Romans 8:26 - Likewise the Spirit also helpeth our infirmities: for we know not what we should pray for as we ought: but the Spirit itself maketh intercession for us with groanings which cannot be uttered.
Philippians 4:19 - But my God shall supply all your need according to his riches in glory by Christ Jesus.
Hebrews 4:15-16 - For we have not an high priest which cannot be touched with the feeling of our infirmities; but was in all points tempted like as [we are, yet] without sin. (Read More...)
Topics and verses are auto-generated from user searches. If a verse or topic does not belong, please contact us. Some scripture references/categories courtesy of Open Bible .info under CC BY 3.0
Related Bible Dictionary Terms:
Search the King James Version (KJV) for more references about Weakness...
Bible Verses About God
Bible Verses About Children
Bible Verses About Divorce


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[PHOTO] iris-photos | Wed, 14 Apr 1999 22:54:05" />
Here is the two I have in bloom. The Southland is a MDB. The purple one
is a old unknow that was given to me. They didn't know the name.
Barney Lou Hillyard in rainie western Kentucky where the temp is
dropping the last I heard it was 54 degrees.


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12 Days Of Beyoncé: "Superpower" Video Fashion
On this last day of "Beyoncé" fever, we bring you the most 'powerful' video, "Superpower".
Directed by Jonas Åkerlund and styled by Bea Åkerlund, the melodic song that features Frank Ocean has its story shown through a fiery revolt. Giving us an apocalyptic theme, we see ripped jeans, leather and distress animal skin everywhere.
While the song is not a favorite, the clothing is interesting because of the renegade theme. Beyoncé wears two looks, the first a camo print jacket (similar style found here), booty shorts– a running theme in all these videos– and fishnet stockings.
For her second look, the songstress shows under boob in a ripped top, canvas chaps and platform boots. This look has a personality of its own. Not organic to Bey's southern sweetie image, she definitely strives to show her edgy side in this video.