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The COllaborative project of Development of Anthropometrical measures in Twins (CODATwins) project is a large international collaborative effort to analyze individual-level phenotype data from twins in multiple cohorts from different environments. The main objective is to study factors that modify genetic and environmental variation of height, body mass index (BMI, kg/m2) and size at birth, and additionally to address other research questions such as long-term consequences of birth size. The project started in 2013 and is open to all twin projects in the world having height and weight measures on twins with information on zygosity. Thus far, 54 twin projects from 24 countries have provided individual-level data. The CODATwins database includes 489,981 twin individuals (228,635 complete twin pairs). Since many twin cohorts have collected longitudinal data, there is a total of 1,049,785 height and weight observations. For many cohorts, we also have information on birth weight and length, own smoking behavior and own or parental education. We found that the heritability estimates of height and BMI systematically changed from infancy to old age. Remarkably, only minor differences in the heritability estimates were found across cultural–geographic regions, measurement time and birth cohort for height and BMI. In addition to genetic epidemiological studies, we looked at associations of height and BMI with education, birth weight and smoking status. Within-family analyses examined differences within same-sex and opposite-sex dizygotic twins in birth size and later development. The CODATwins project demonstrates the feasibility and value of international collaboration to address gene-by-exposure interactions that require large sample sizes and address the effects of different exposures across time, geographical regions and socioeconomic status.
Laser interaction with an ultra-thin pre-structured target is investigated with the help of both two-dimensional and three-dimensional particle-in-cell simulations. With the existence of a periodic structure on the target surface, the laser seems to penetrate through the target at its fundamental frequency even if the plasma density of the target is much higher than the laser’s relativistically critical density. The particle-in-cell simulations show that the transmitted laser energy behind the pre-structured target is increased by about two orders of magnitude compared to that behind the flat target. Theoretical analyses show that the transmitted energy behind the pre-structured target is actually re-emitted by electron ‘islands’ formed by the surface plasma waves on the target surfaces. In other words, the radiation with the fundamental frequency is actually ‘surface emission’ on the target rear surface. Besides the intensity of the component with the fundamental frequency, the intensity of the high-order harmonics behind the pre-structured target is also much enhanced compared to that behind the flat target. The enhancement of the high-order harmonics is also related to the surface plasma waves generated on the target surfaces.
Global inequity in access to and availability of essential mental health services is well recognized. The mental health treatment gap is approximately 50% in all countries, with up to 90% of people in the lowest-income countries lacking access to required mental health services. Increased investment in global mental health (GMH) has increased innovation in mental health service delivery in LMICs. Situational analyses in areas where mental health services and systems are poorly developed and resourced are essential when planning for research and implementation, however, little guidance is available to inform methodological approaches to conducting these types of studies. This scoping review provides an analysis of methodological approaches to situational analysis in GMH, including an assessment of the extent to which situational analyses include equity in study designs. It is intended as a resource that identifies current gaps and areas for future development in GMH. Formative research, including situational analysis, is an essential first step in conducting robust implementation research, an essential area of study in GMH that will help to promote improved availability of, access to and reach of mental health services for people living with mental illness in low- and middle-income countries (LMICs). While strong leadership in this field exists, there remain significant opportunities for enhanced research representing different LMICs and regions.
Triploid and pentaploid breeding is of great importance in agricultural production, but it is not always easy to obtain double ploidy parents. However, in fishes, chromosome ploidy is diversiform, which may provide natural parental resources for triploid and pentaploid breeding. Both tetraploid and hexaploid exist in Schizothorax fishes, which were thought to belong to different subfamilies with tetraploid Percocypris fishes in morphology, but they are sister genera in molecule. Fortunately, the pentaploid hybrid fishes have been successfully obtained by hybridization of Schizothorax wangchiachii (♀, 2n = 6X = 148) × Percocypris pingi (♂, 2n = 4X = 98). To understand the genetic and morphological difference among the hybrid fishes and their parents, four methods were used in this study: morphology, karyotype, red blood cell (RBC) DNA content determination and inter-simple sequence repeat (ISSR). In morphology, the hybrid fishes were steady, and between their parents with no obvious preference. The chromosome numbers of P. pingi have been reported as 2n = 4X = 98. In this study, the karyotype of S. wangchiachii was 2n = 6X = 148 = 36m + 34sm + 12st + 66t, while that the hybrid fishes was 2n = 5X = 123 = 39m + 28sm + 5st + 51t. Similarly, the RBC DNA content of the hybrid fishes was intermediate among their parents. In ISSR, the within-group genetic diversity of hybrid fishes was higher than that of their parents. Moreover, the genetic distance of hybrid fishes between P. pingi and S.wangchiachii was closely related to that of their parental ploidy, suggesting that parental genetic material stably coexisted in the hybrid fishes. This is the first report to show a stable pentaploid F1 hybrids produced by hybridization of a hexaploid and a tetraploid in aquaculture.
This study aims to investigate the prevalence and genotype distribution of anal human papillomavirus (HPV) infection among men with different sexual orientations with or without human immunodeficiency virus (HIV) in China. A cross-sectional study was conducted during 2016–2017 in Taizhou City, Zhejiang Province. Convenient sampling was used to recruit male participants from HIV voluntary counselling and testing clinics and Center for Disease Control and Prevention. A face-to-face questionnaire interview was administered and an anal-canal swab was collected for HPV genotyping. A total of 160 HIV-positive and 113 HIV-negative men participated in the study. The prevalence of any type HPV was 30.6% for heterosexual men, 74.1% for homosexual and 63.6% for bisexual men among HIV-positive participants, while the prevalence was 8.3%, 29.2% and 23.8% respectively among HIV-negatives. The most prevalent genotypes were HPV-58 (16.9%), HPV-6 (15.6%) and HPV-11 (15.0%) among HIV-positive men, and were HPV-16 (4.4%), HPV-52 (4.4%) and HPV-6 (3.5%) among HIV-negative men. Having ever had haemorrhoids and having ever seen blood on tissue after defaecation was associated with HPV infection. One-fourth of the HPV infections in this study population can be covered by the quadrivalent vaccine in market. The highly prevalent anal HPV infection among men especially HIV-infected men calls for close observation and further investigation for anal cancer prevention.
The seasonality of individual influenza subtypes/lineages and the association of influenza epidemics with meteorological factors in the tropics/subtropics have not been well understood. The impact of the 2009 H1N1 pandemic on the prevalence of seasonal influenza virus remains to be explored. Using wavelet analysis, the periodicities of A/H3N2, seasonal A/H1N1, A/H1N1pdm09, Victoria and Yamagata were identified, respectively, in Panzhihua during 2006–2015. As a subtropical city in southwestern China, Panzhihua is the first industrial city in the upper reaches of the Yangtze River. The relationship between influenza epidemics and local climatic variables was examined based on regression models. The temporal distribution of influenza subtypes/lineages during the pre-pandemic (2006–2009), pandemic (2009) and post-pandemic (2010–2015) years was described and compared. A total of 6892 respiratory specimens were collected and 737 influenza viruses were isolated. A/H3N2 showed an annual cycle with a peak in summer–autumn, while A/H1N1pdm09, Victoria and Yamagata exhibited an annual cycle with a peak in winter–spring. Regression analyses demonstrated that relative humidity was positively associated with A/H3N2 activity while negatively associated with Victoria activity. Higher prevalence of A/H1N1pdm09 and Yamagata was driven by lower absolute humidity. The role of weather conditions in regulating influenza epidemics could be complicated since the diverse viral transmission modes and mechanism. Differences in seasonality and different associations with meteorological factors by influenza subtypes/lineages should be considered in epidemiological studies in the tropics/subtropics. The development of subtype- and lineage-specific prevention and control measures is of significant importance.
Distillers dried grains with solubles (DDGS) are highly susceptible to lipid oxidation because DDGS contain about 10% crude fat, which is largely composed of polyunsaturated fatty acids. l-carnitine serves an important function in fatty acids β-oxidation, and also has antioxidant properties. The objective of this study was to examine the effects of l-carnitine in the DDGS diet of gestating and lactating sows on reproductive performance, milk composition and antioxidant status of sows and their offspring. One hundred and twenty sows (Landrace×Large white, mean parity 4.2, initial BW 230 kg) were randomly allotted to 1 of 4 dietary treatments (n=30 sows/treatment). Treatments were arranged as a 2×2 factorial with two levels of dietary DDGS (0 v. 250 g/kg in gestating diets and 400 g/kg in lactating diets) and two levels of dietary l-carnitine (0 v. 100 mg/kg in gestating diets and 0 v. 200 mg/kg in lactating diets). Distillers dried grains with solubles had no significant effect on litter size but significantly reduced the birth weights and weaning weights of piglets (P<0.05). Distillers dried grains with solubles reduced the antioxidant enzyme activities (P<0.05) and increased the malondialdehyde level in the plasma of sows on day 60 of gestation (P=0.004) and day 14 of lactation (P=0.008). The compositions of colostrum and milk were not affected by inclusion of DDGS and dietary l-carnitine (P>0.05). Supplementing the diets with l-carnitine had no significant effect of total litter size (P>0.05) but increased the number of piglets born alive and piglets weaned, birth weight and weaning weight of piglets and litter weight at birth and weaning (P<0.05). l-carnitine supplementation also increased the concentration of l-carnitine in milk and l-carnitine status of piglets (P<0.05). The antioxidant enzyme activities of new born and weaning piglets were increased (P<0.05) by maternal dietary l-carnitine but this did not extend to finishing pigs. In conclusion, including DDGS in the sows diet could induce oxidative stress, which may be associated with the reduced individual birth and weaning weight of piglets. Dietary l-carnitine supplementation improved the antioxidant and l-carnitine status of sows, which may be associated with the improved reproduction and piglet performance and the antioxidant status of piglets at birth and weaning. There were no interactions between DDGS and l-carnitine.
High ambient temperatures affect animal production and welfare in tropical and sub-tropical regions of the world. Feed intake, growth rate, mortality, egg production, hatchability and other production traits related to the economic success of the poultry industry are adversely affected by severe heat stress. In general, heat stress induces the activity of the neuroendocrine system, resulting in activation of the hypothalamic-pituitary-adrenal (HPA) axis, and elevated corticosterone (CORT) concentrations, which affects metabolism and immune responses. These include negative regulation of metabolic hormones, antibody production and heterophil to lymphocyte (H/L) ratio. Heat stress increases mitochondrial activity, causing reactive species overproduction which disrupts the antioxidant balance, leading to oxidative stress damage of membranes, protein and DNA. Heat stress stimulates the central nervous system (CNS), which significantly reduces daily gain, feed intake and FCR in poultry. Consequently, from an animal husbandry perspective, intervention strategies to relieve heat stress conditions have been the focus of many published studies. This review describes the effect of high temperature on production, behavioural, biochemical and immune responses, including oxidative damage that occur during heat stress in poultry, in broilers and laying hens. Moreover, nutritional interventions to alleviate the negative consequence of heat stress is discussed.
Hydrogen is an important intermediate that is produced during carbohydrate fermentation to volatile fatty acid and utilized by methanogens to produce methane in the rumen. Ruminal volatile fatty acid and dissolved methane concentrations are more than 500 times greater than dissolved hydrogen concentration. Therefore, we hypothesized that dissolved hydrogen might have a higher sensitivity in response to dietary changes compared with volatile fatty acid and dissolved methane. Using goats, we investigated the effects of increasing dietary starch content (maize replaced with wheat bran) and supplementing with rhubarb rhizomes and roots on the relationships among dissolved hydrogen, dissolved methane and other fermentation end products. The study was conducted in a replicated 4×4 Latin square with a 2×2 factorial arrangement of four treatments: two starch levels (220 v. 320 g/kg dry matter (DM)), without and with rhubarb supplement (0% v. 2.8% of total mixed ration). Increased dietary starch and rhubarb supplementation did not alter volatile fatty acid concentrations or methane emissions in terms of g/day, g/g DM intake and g/g organic matter digested. However, goats fed the high-starch diet had greater dissolved hydrogen (P=0.005) and relative abundance of Selenomonas ruminantium (P<0.01), and lower (P=0.02) copy number of protozoa than those fed the low-starch diet. Rhubarb increased ruminal dissolved H2 (P=0.03) and total volatile fatty acid concentration (P<0.001), but decreased copies of bacteria (P=0.002). In conclusion, dissolved hydrogen appears to be more sensitive to dietary changes with starch content and rhubarb supplementation, when compared with volatile fatty acid concentrations and methane production.
Analysis of the potato yield gap and the corresponding nutrient gap can help in devising strategies and measurements to increase productivity for closing the gaps through improved practices. On-farm experiments conducted in the main potato production areas of northwest China were used to determine attainable yield. Official statistical data were used to determine the actual on-farm yield. Yield gap was the difference between attainable yield and actual on-farm yield. Nutrient gap was calculated by dividing the size of yield gap by partial factor productivity. Results indicated that nitrogen (N), phosphorus (P) and potassium (K) fertilization increased potato yield by an average of 1169–7625, 2937–5336 and 2331–7338 kg/ha, respectively. The maximum attainable yields (the 90th percentile yields) were 50 145, 37 855, 30 261 and 56 616 kg/ha and the average actual on-farm yield were 14 179, 16 732, 10 271 and 19 990 kg/ha in the Inner Mongolia Autonomous Region (IMAR), Gansu, Ningxia and Qinghai provinces, respectively. In the above four regions, yield would need to increase by 165, 70, 112 and 121% from actual yield to reach 75% of attainable yield. Compared with recent 3-year average NPK rates by farmers, the total NPK rates need to increase by 90.1–134.3% for IMAR, 42.9–69.2% for Gansu, 68.1–111.2% for Ningxia and 48.1–83.8% for Qinghai to improve productivity to near the 75% attainable yield. In conclusion, the high yield responses to fertilizer application provide opportunities to close the large yield gaps through balanced nutrition.
Cultivated pastures in southern China are being used to improve forage productivity and animal performance, but studies on grazing behaviour of goats in these cultivated pastures are still rare. In the current study, the grazing behaviour of Yunling black goats under low (5 goats/ha) and high (15 goats/ha) stocking rates (SRs) was evaluated. Data showed that the proportion of time goats spent on activities was: eating (0.59–0.87), ruminating (0.05–0.35), walking (0.03–0.06) and resting (0.01–0.03). Compared with low SR, goats spent more time eating and walking, and less time ruminating and resting under high SR. Goats had similar diet preferences under both SR and preferred to eat grasses (ryegrass and cocksfoot) more than a legume (white clover). The distribution of eating time on each forage species was more uniform under high v. low SR. Bites/step, bite weight and daily intake were greater under low than high SR. Results suggest that the SR affects grazing behaviour of goats on cultivated pasture, and identifying an optimal SR is critical for increasing bite weight and intake.
Direct numerical simulation of intense laser–solid interactions is still of great challenges, because of the many coupled atomic and plasma processes, such as ionization dynamics, collision among charged particles and collective electromagnetic fields, to name just a few. Here, we develop a new particle-in-cell (PIC) simulation code, which enables us to calculate laser–solid interactions in a more realistic way. This code is able to cover almost ‘all’ the coupled physical processes. As an application of the new code, the generation and transport of energetic electrons in front of and within the solid target when irradiated by intense laser beams are studied. For the considered case, in which laser intensity is
and pre-plasma scale length in front of the solid is
, several quantitative conclusions are drawn: (i) the collisional damping (although it is very weak) can significantly affect the energetic electrons generation in front of the target, (ii) the Bremsstrahlung radiation will be enhanced by 2–3 times when the solid is dramatically heated and ionized, (iii) the ‘cut-off’ electron energy is lowered by an amount of 25% when both collision damping and Bremsstrahlung radiations are included, and (iv) the resistive electromagnetic fields due to Ohmic heating play nonignorable roles and must be taken into account in such interactions.
The gastrointestinal tract (GIT) of animals is capable of sensing various kinds of nutrients via G-protein coupled receptor-mediated signaling transduction pathways, and the process is known as ‘gut nutrient chemosensing’. GPR40, GPR41, GPR43 and GPR119 are chemoreceptors for free fatty acids (FFAs) and lipid derivatives, but they are not well studied in small ruminants. The objective of this study is to determine the expression of GPR40, GPR41, GPR43 and GPR119 along the GIT of kid goats under supplemental feeding (S) v. grazing (G) during early development. In total, 44 kid goats (initial weight 1.35±0.12 kg) were slaughtered for sampling (rumen, abomasum, duodenum, jejunum, ileum, cecum, colon and rectum) between days 0 and 70. The expression of GPR41 and GPR43 were measured at both mRNA and protein levels, whereas GPR40 and GPR119 were assayed at protein level only. The effects of age and feeding system on their expression were variable depending upon GIT segments, chemoreceptors and expression level (mRNA or protein), and sometimes feeding system × age interactions (P<0.05) were observed. Supplemental feeding enhanced expression of GPR40, GPR41 and GPR43 in most segments of the GIT of goats, whereas G enhanced expression of GPR119. GPR41 and GPR43 were mainly expressed in rumen, abomasum and cecum, with different responses to age and feeding system. GPR41 and GPR43 expression in abomasum at mRNA level was greatly (P<0.01) affected by both age and feeding system; whereas their expression in rumen and abomasum at protein level were different, feeding system greatly (P<0.05) affected GPR41 expression, but had no effect (P>0.05) on GPR43 expression; and there were no feeding system×age interactions (P>0.05) on GPR41 and GPR43 protein expression. The expression of GPR41 and GPR43 in rumen and abomasum linearly (P<0.01) increased with increasing age (from days 0 to 70). Meanwhile, age was the main factor affecting GPR40 expression throughout the GIT. These outcomes indicate that age and feeding system are the two factors affecting chemoreceptors for FFAs and lipid derivatives expression in the GIT of kids goats, and S enhanced the expression of chemoreceptors for FFAs, whereas G gave rise to greater expression of chemoreceptors for lipid derivatives. Our results suggest that enhanced expression of chemoreceptors for FFAs might be one of the benefits of early supplemental feeding offered to young ruminants during early development.
Moringa oleifera seeds are currently being used as a livestock feed across tropical regions of the world due to its availability and palatability. However, limited knowledge exists on the effects of the raw seeds on ruminant metabolism. As such, the rumen stimulation technique was used to evaluate the effects of substituting increasing concentrations of ground Moringa seeds (0, 100, 200 and 400 g/kg concentrate dry matter (DM)) in the diet on rumen fermentation and methane production. Two identical, Rusitec apparatuses, each with eight fermenters were used with the first 8 days used for adaptation and days 9 to 16 used for measurements. Fermenters were fed a total mixed ration with Urochloa brizantha as the forage. Disappearance of DM, CP, NDF and ADF linearly decreased (P<0.01) with increasing concentrations of Moringa seeds in the diet. Total volatile fatty acid production and the acetate to propionate ratio were also linearly decreased (P<0.01). However, only the 400 g/kg (concentrate DM basis) treatment differed (P<0.01) from the control. Methane production (%), total microbial incorporation of 15N and total production of microbial N linearly decreased (P<0.01) as the inclusion of Moringa seeds increased. Though the inclusion of Moringa seeds in the diet decreased CH4 production, this arose from an unfavourable decrease in diet digestibility and rumen fermentation parameters.
Toca 511 (vocimagene amiretrorepvec) is an investigational retroviral replicating vector that selectively infects dividing cancer cells, integrates into the genome and replicates due to immune defects in tumors. Toca 511 spreads through tumors and stably delivers the gene encoding an optimized yeast cytosine deaminase that converts the prodrug Toca FC (investigational, extended-release of 5-fluorocytosine) into 5-fluorouracil. In preclinical models, 5-fluorouracil kills infected dividing cancer cells, myeloid derived suppressor cells and tumor associated macrophages, enabling immune activation against the tumor. In this dose ascending Ph1 trial (NCT01470794), Toca 511 was injected into the resection cavity wall of patients with rHGG, followed by courses of oral Toca FC. Additional cohorts included combination with bevacizumab or lomustine. Across the Ph1 program, the safety profile remains favorable. Objective responses (ORs) were assessed by IRR using MRI scans prior to Toca FC treatment as baseline. ORs occurred 6-19 months after Toca 511 administration, suggesting an immunologic mechanism. The ORs were observed in 4 patients with IDH1-wildtype and 2 patients with IDH1-mutant tumors, including 5 complete responses (CRs) with the investigational therapy alone, and 1 CR in combination with bevacizumab. The median duration of response (mDoR) was 35.1+ months. As of AUG2017, all responders were CR and remain alive. In a 23-patient subgroup who received high doses of Toca 511 and met Ph3 trial criteria, mOS was 14.4 months, 3-year survival rate was 26.1%, and mDoR was 35.7+ months with a durable response rate of 21.7%. Data suggest a positive association of durable response with OS.
The increasing tropospheric ozone concentration [O3] strongly affects plant growth. However, the response of belowground processes in rice (Oryza sativa L.) systems to higher O3 is not well understood. The grain production, belowground biomass partitioning, root morphology and activity of rice (cv. Shanyou 63) were investigated in a free-air O3 enrichment platform at four key growth stages. Elevated O3 (EO3, 50% above the ambient O3) significantly decreased the grain yield and total biomass at the grain milky mature stage, root biomass at the tillering stage and root to shoot ratios (RRS) at the flowering and grain filling stages. The effects of EO3 on root morphology and activity varied among rice growth stage. EO3 significantly decreased root length, density, area, diameter and volume at the flowering stage, but EO3 significantly decreased various root morphological indices at the tillering, grain filling and milky mature stages. EO3 significantly increased the specific root respiration rate (root activity) and root respiration rate (autotrophic respiration) at grain filling and milky mature stages. Higher root autotrophic respiration and lower RRS in response to EO3 would reduce allocation of assimilated carbon to root growth, adversely affecting rice productivity. Our findings are critical for understanding the O3-induced impairment of belowground processes and carbon cycling in rice cropping systems and breeding of O3-tolerant cultivars under higher [O3] scenarios.
Many shorebird populations are in decline along the East Asian-Australasian Flyway. The rapid loss of coastal wetlands in the Yellow Sea, which provide critical stop-over sites during migration, is believed to be the cause of the alarming trends. The Yalu Jiang coastal wetland, a protected area in the north Yellow Sea, supports the largest known migratory staging populations of Bar-tailed Godwits Limosa lapponica (menzbieri and baueri subspecies) and Great Knots Calidris tenuirostris. Monitoring of the macrozoobenthos food for these shorebirds from 2011 to 2016 showed declines of over 99% in the densities of the bivalve Potamocorbula laevis, the major food here for both Bar-tailed Godwits and Great Knots. The loss of the bivalve might be caused by any combination of, but not limited to: (1) change in hydrological conditions and sediment composition due to nearby port construction, (2) run-off of agrochemicals from the extensive shoreline sea cucumber farms, and (3) parasitic infection. Surprisingly, the numbers of birds using the Yalu Jiang coastal wetland remained stable during the study period, except for the subspecies of Bar-tailed Godwit L. l. menzbieri, which exhibited a 91% decline in peak numbers. The lack of an overall decline in the number of bird days in Great Knots and in the peak numbers of L. l. baueri, also given the published simultaneous decreases in their annual survival, implies a lack of alternative habitats that birds could relocate to. This study highlights that food declines at staging sites could be an overlooked but important factor causing population declines of shorebirds along the Flyway. Maintaining the quality of protected staging sites is as important in shorebird conservation as is the safeguarding of staging sites from land claim. Meanwhile, it calls for immediate action to restore the food base for these beleaguered migrant shorebirds at Yalu Jiang coastal wetland.
Litter size has a great impact on the profit of swine producers. Uterine development is an important determinant of reproduction efficiency and could hence affect litter size. Chinese Erhualian pig is one of the most prolific breeds in the world, even though large phenotypic variation in litter size was observed within Erhualian sows. To dissect the genetic basis of the phenotypic variation, we herein conducted genome-wide association studies for total number born and number born alive (NBA) of Erhualian sows. In total, one significant single nucleotide polymorphism (SNP) (P<1.78e−06) and 11 suggestive SNPs (P<3.57e−05) were identified on 10 chromosomes, confirming seven previously reported quantitative trait loci (QTL) and uncovering six QTL for litter size or uterus length. One locus on Sus scrofa chromosome (SSC) 13 (79.28 to 90.43 Mb) harbored a cluster of suggestive SNPs associated with multiparous NBA. The SNP (rs81447100) within this region was confirmed to be significantly (P<0.05) associated with litter size in Erhualian (n=313), Sutai (n=173) and Yorkshire (n=488) populations. Retinol binding protein 2 and retinol binding protein 1 functionally related to the development of uterus were located in a region of 2 Mb around rs81447100. Moreover, four genes related to embryo implantation and development were also detected around other significant SNPs. Taken together, our findings provide a potential marker (rs81447100) for the genetic improvement of litter size not only in Chinese Erhualian pigs but also in European commercial pig breeds like Yorkshire, and would facilitate the final identification of causative variant(s) underlying the effect of SSC13 QTL on litter size.