본 연구에서는 효과적이고 안전한 화장품 성분 탐색을 위해 항산화 활성 가이드라인에 따라 다양한 식물 추출물을 대상으로 효능을 확인하였으며 이 중 가장 효능이 좋은 식물을 발굴하고 이를 최적의 비율로 혼합하여 Charmzone Extract (CZE)로 명명하였다. CZE는 하엽(Nelumbo nucifera leaves), 삼백초(Saururus chinensis) 및 와송(Orostachys japonica)으로 구성된 식물복합추출물로, 본 연구에서는 CZE의 항노화 화장품 원료로서의 개발 가능성을 확인해 보고자 하였다. CZE는 자유 라디칼 소거 활성을 가지고 있었으며, 인간 각질 형성세포주인 HaCaT 및 인간 섬유아세포 NHDF에서 활성산소종의 생성을 억제하고 총항산화능을 증가시켰다. 그리고 NHDF에서 제1형 프로콜라겐의 분비를 촉진하였고, B16F10 세포에서 멜라닌의 생성을 감소시켰으며, RAW264.7 세포에서 lipopolysaccharide 자극에 의한 산화질소의 생성을 농도 의존적으로 저해하였다. 또한, CZE는 HaCaT의 이동을 촉진하는 효능을 나타내었다. 결과를 종합하면, CZE는 항산화, 주름개선, 미백, 항염증 및 상처치유의 다효능을 나타내며, 이러한 효능을 통해 피부 기능을 향상시킬 수 있을 것이라고 기대된다. 그러 므로 식물복합추출물 CZE는 피부 항노화용 화장품 성분으로 응용 가능하다.

지금까지 저점도 메이크업 유중수형 에멀전을 함침재에 침적시켜 휴대 간편성과 사용 편리성을 높인 화 장품이 개발되어 왔다. 종래에는 주로 경도가 높고 다공의 망상구조를 갖는 폴리우레탄 스펀지가 함침재로서 사용되어 왔으며, 이러한 스펀지는 구조적인 특성상 화장료 함침이 잘 되기 때문에 제조가 용이하다는 장점을 가지고 있지만, 초기 토출량이 높고, 사용 횟수에 따라 토출량의 감소가 큰 특징을 가지고 있었다. 본 연구에서는 이러한 문제를 해결할 수 있는 새로운 발포형태의 스펀지를 개발하고 이를 활용한 다양한 특징을 갖는 메이크업 유중수형 에멀전에 관하여 연구하고자 한다. 우선 상기 목적에 맞는 다양한 소재를 조사하고, 이를 토대로 natural rubber (NR)와 styrene butadiene rubber (SBR)의 조합을 통해 성형성이 우수하며 화장료 조성물의 적절한 흡수 및 배출 능력을 가지는 새로운 물질을 개발하였다. 이러한 함침재는 고무와 같은 탄성성질의 부드러 움과 높은 신율 및 균일한 토출력을 가지는 특징을 확인할 수 있었다. 화장료 조성물로 사용되는 여러 종류의 오일의 극성도(polarity)에 따른 신규 스펀지의 특성을 이해하고, 메이크업 유중수형 에멀전의 점성을 조절하여 메이크업 제품 개발이 가능함을 확인하였으며, 향후 새로운 특성을 갖는 화장료 함침재로써의 가치가 매우 높을 것으로 기대된다.

한중 양국은 오랜 역사기간동안 문화와 정치, 종교와 예술에서 끊임없는 교류를 통하여 우리나라 는 자체적인 우수문화를 창달하였다. 중국의 선진문물과 문화를 도입하여 발전시키고 이를 이웃국가 인 일본에 전파하는 중간 매개체 역할도 수행해왔다. 특히 인쇄와 종이제조 기술의 유입과 발달은 중 국의 문학작품을 수용하여 한국고유의 작품탄생이 탄생할 수 있는 중요한 토대가 되었다. 역사적으로 한국에 유입된 중국 고대문학 작품 수는 총 483종이며 이중 전등신화와 전등여화를 포 함한 소설이 464종, 원나라 잡극 서상기를 포함한 희곡 19종이다. 특히 조선왕 연산군이 직접 하교하 여 중국 서적을 구매해서 들여오라는 기록이 있다. 조선왕조실록 연산군일기에 중국에 파견하는 사절 단에게 명나라에서 서적을 구매해 올 것을 명하였다. 이를 통하여 우리는 파견 사절단이 서적 유입의 중요한 역할을 담당했음을 알 수 있다. 이렇게 유입된 서적 중 전등신화와 전등이화는 조선 초기 한국의 최초 소설인 금오신화에 커다란 영향을 주었다. 이후 전기소설집 기재기이의 탄생은 17세기 몽유록 소설의 대표작인 구운몽의 탄생에 중요한 가교 역할을 하였다. 또한 대표적인 애정소설 운영전과 주생전 그리고 최초의 한글소설 홍길 동전 탄생에 지대한 영향을 주었다. 무역에 의한 중국 서적 유입은 한국의 고전소설의 탄생과 발전에 밀접한 관련이 있음을 알 수 있다. 현대사회에서 문화컨텐츠는 고전소설을 중요한 소재로 활용하고 있다. 이러한 문화산업을 활용하 여 수출과 관광자원으로 활용한다면 국가 경제 이익창출에 크게 기여할 수 있을 것이다. 미래에 문화 산업 수출의 중요성은 더욱더 강조될 것이다. 오래전에 국내로 들여온 무역서적 중 전등이종의 국내 유입은 우리의 자체적인 문학작품탄생의 모티브가 되었으며 독자적인 소설 발전에 크게 기여한 것이 다. 이후 우리는 우수한 고전문학작품을 문화컨텐츠 소재로 잘 활용한다면 문화산업발전과 나아가 세 계 무역교류를 이끄는 선두 주자가 될 것이다. 따라서 본 논문을 통하여 중국서적의 무역유입과 현 재의 한국 경제에 깊은 영향관계를 밝히고자 한다.

Sprouty (Spry) genes encode inhibitors of the receptor tyrosine kinase signaling cascade, which plays important roles in stem cells. However, the role of Spry4 in the stemness of embryonic stem cells has not been fully elucidated. Here, we used mouse embryonic stem cells (mESCs) as a model system to investigate the role of Spry4 in the stem cells. Suppression of Spry4 expression results in the decreases of cell proliferation, EB formation and stemness marker expression, suggesting that Spry4 activity is associated with stemness of mESCs. Teratoma assay showed that the cartilage maturation was facilitated in Spry4 knocked down mESCs. Our results suggest that Spry4 is an important regulator of the stemness and differentiation of mESCs.

Although the toxicological impacts of the xenoestrogen bisphenol-A (BPA) have been studied extensively, but its mechanism of action is poorly understood. Eventually, no standard method exists for evaluating the possible health hazard of BPA. Considering mice spermatozoa as a potential in vitro model, here we demonstrated the effects of BPA exposure (0.0001, 0.01, 1, and 100 μM for 6 h) on spermatozoa and the related mechanisms of action. Our results demonstrated that high concentrations of BPA negatively affect sperm motility, viability, intracellular ATP, and mitochondrial functions by activating the mitogen-activated protein kinase, phosphatidylinositol 3-kinase, and protein kinase-A pathways. The same doses were also employed to identify the differential expressed proteins of exposure and screen their functional affiliation to diseases using sperm proteomics and informatics, respectively. Our results demonstrated that a high concentration of BPA (100 μM) induced differential expression (> 2-fold) of 24 proteins in spermatozoa (16 down- and 9 up-regulated), that are putatively involved in the pathogenesis of several diseases. To the best of our knowledge, this is the first study to demonstrate the mechanisms of BPA action in spermatozoa and to identify the possible biomarkers of exposure. Moreover, we anticipated that current strategy might apply for the hazard assessment of other toxicological agents.

Future space missions will have larger telescopes in order to look deeper into space while improvising on spatial resolution. The primary mirrors for these telescopes will be so large that using a monolithic mirror will be nearly impossible because of the difficulties associated with its fabrication, transportation, and installation on a launch vehicle. The feasibility of launching these huge mirrors is limited because of their small launch fairing diameter. The aerodynamic shape of the fairing requires a small diameter, but the height of the launch vehicle, which is available for designers to utilize, is larger than the fairing diameter. This paper presents the development of an axial deployment mechanism based on the screw jack principle. The mechanism was designed and developed, and a prototype was constructed in order to demonstrate a lab model.

The wild relatives of soybean [Glycine soja Sieb. and Zucc.] have curly/wavy nature whereas cultivated varieties are upright. Such morphological characteristics have agronomic importance too. To investigate the molecular mechanism of development contributing to coiled morphology, screening was carried out to look for Arabidopsis mutants in activation tagging lines obtained by activation T-DNA treatment that have curly/wavy morphology. A mutant named Coiled Branch 1 (cbr1), is found to have a wavy and curly morphology with coiling branches. Plasmid rescue and genomic southern blot analysis revealed the site of T-DNA insertion in the genome. RT-PCR was performed to monitor expression levels of the genes adjacent to the T-DNA integration sites, and showed the activation of an E3 ubiquitin ligase gene. Database search showed that the gene with the RING domain belongs to a family of E3 ubiquitin ligases. Complementation test by overexpression and RNA interference of the gene was also carried out. The complementation test results showed that the novel gene activation tagging affected the cbr1 mutant phenotypes. Ubiquitylation has been linked virtually to every cellular process including plant development. E3 ubiquitin ligase has been reported to recognize target proteins that are to be ubiquinated for further degradation by the proteasome complex. Further, more detailed studies are needed to identify the specific substrate(s) of the novel E3 ubiquitin ligase gene.

Soybean [Glycine max (L.) Merr.] seeds are abundant in high-quality proteins and fats. In addition, soybean seeds are also rich in secondary metabolites, such as isoflavones, lecithin, and saponins. Triterpene saponins are major components of these physiologically active metabolites in soybean seeds. Soybean saponins are classified as group A and DDMP saponins. Among them group A saponins are undesirable component of food products due to bitterness and astringency and also cause foaming in tofu production. Whereas, DDMP saponins and their derivatives are less bitter and astringent and beneficial to human health when consumed as regular diet. Therefore, reducing the group A saponins or increasing the DDMP saponins are required to improve the food quality. The present study focused to identify and characterize the gene which is encoding a protein responsible for biosynthesis of DDMP saponins. EMS mutant lines (sg-7-1 & sg-7-2) which lack DDMP saponins were developed. The breeding cross has been made with these two mutants with two cultivars, Pungsannamul and Wooram to study the segregation and genetic linkage analysis, respectively. The segregation analysis showed that the mutant phenotype is controlled by single recessive gene. TLC analysis for phenotyping F2 population of Wooram X sg-7-1 showed mutant, wild and heterozygous types. To surprise two more patterns were detected and they were named as strange type1 (ST1) and strange type2 (ST2). Further, SSR marker analysis will be carried out to locate the gene which encoding a protein responsible for biosynthesis of DDMP saponins.

Blueberry (Vaccinium spp.) is a member of the Ericaceae and eleven varieties have been registered at the Korea Seed & Variety Service for Plant Variety Protection (PVP). This study was to develop simple sequence repeat (SSR) markers next generation sequencing (NGS) analysis and to analysis genetic relationship of blueberry 31 varieties. Highbush blueberry ‘Camellia’ and rabbiteye blueberry ‘Alapaha’ varieties were used as sequencing materials. Out of total 987 SSR primers detected between ‘Camellia’ and ‘Alapaha’, 148 SSR primers were initially applied to select SSR markers for identification of blueberry varieties. Fourteen SSR markers showed polymorphism between 8 varieties. Seven SSR markers showed reproducibility and clear peak among 14 SSR markers. Genetic relationships of 31 blueberry varieties were analyzed and identified using 7 SSR markers. A total of 30 polymorphic SSR alleles were obtained and two to seven alleles were detected for each locus with an average of 4.3 alleles per locus. Average polymorphism information content was 0.556, ranging from 0.374 to 0.714. Genetic distance of clusters ranged from 0.38 to 0.93 by unweighted pair-group method with arithmetical average based on Jaccard’s distance coefficients. These newly developed SSR markers indicate usefulness for variety identification related to seed dispute and distinctness, uniformity and stability (DUS) test for blueberry.

Small RNAs, such as microRNAs (miRNAs) and small interfering RNAs (siRNAs), play crucial roles in post-transcriptional gene silencing (PTGS) in eukaryotes. Small RNAs function cell-autonomously as well as non-cell-autonomously. It has been well characterized that pathogenic fungi secrete some effector molecules facilitating their infection into plants. However, it is unclear whether molecules produced in plant cells are able to move into fungal cells during infection. To test if small RNAs generated from plant cells can move to fungal cells during infection, we generated transgenic Arabidopsis and rice plants expressing siRNAs targeting GFP gene generated from double-stranded RNA interference (dsRNAi) constructs for GFP gene. And then these transgenic plants were inoculated with transgenic rice blast fungus, Magnaporthe oryzae, expressing GFP transgene. Here, we showed that ectopic expression of siRNAs targeting GFP gene in transgenic plants significantly suppressed GFP expression in rice blast fungi inoculated, indicating that small RNA molecules generated in plant cells can move into infected fungal cells and efficiently degrade fungal GFP transcripts. Our results would provide a new small RNA-based strategy for the development of resistant crops against fungal pathogens.

Aroma in rice (Oryza sativa L.) results from the loss of function of the betaine aldehyde dehydrogenase (Badh2) gene on chromosome 8. An 8-bp deletion in exon 7 of Badh2 was reported to be the main allele functionally associated with fragrance. The discovery of new functional alleles will provide additional genetic resources to improve fragrant rice. In this study, we sequenced the Badh2 gene in 30 rice accessions and filtered the Badh2 polymorphisms from whole genome re-sequence data of 295 rice accessions. Seven alleles were detected from the sequence data. Six of the seven were known alleles and one was a novel allele (badh2-E12). The novel allele was a 3-bp deletion in exon 12. Five functional markers, targeting six of the seven alleles, were identified. Fourteen accessions were selected to test the utility of these markers. The five molecular markers reliably distinguishing this fragrant rice from other fragrant or non-fragrant rice accessions. Analysis of two F2 rice population validated the genetic markers FME12-3 and FME14I as functional markers. These two markers co-segregated with the fragrance phenotype. These markers will be used in a Badh2 diversity study and to breed improved fragrant rice accessions via marker-assisted selection.

To identify novel signaling components involved in regulation of plant responses to phosphate (Pi) starvation, we screened an Arabidopsis T-DNA activation tagging library for mutants with altered Pi-starvation responses. Here, we report the identification and characterization of novel activation-tagged mutant involved in Pi starvation signaling in Arabidopsis. The hpd (hypersensitive to Pi deficiency) mutant exhibits enhanced phosphate uptake and altered root architectural change under Pi starvation compared to wild type. Expression analysis of auxin-responsive DR5::GUS reporter gene in hpd mutant indicated that both auxin biosynthesis and auxin translocation under Pi starvation are suppressed in hpd mutant plants. Impaired auxin translocation in roots of hpd mutant was attributable to abnormal root architecture changes in Pi starvation conditions. Mis-regulation of auxin translocation in hpd mutant was further confirmed by analysis of expression patterns of auxin efflux carrier proteins, PIN-FORMED (PIN) 1, 2, and 3 fused with GFP. Not only expression levels but also expression domains of PIN proteins were altered in hpd mutant in response to Pi starvation. Molecular genetic analysis of hpd mutant revealed that the mutant phenotype is caused by the lesion in ENHANCED SILENCING PHENOTYPE4 (ESP4) gene whose function is proposed in mRNA 3’-end processing. The results propose that mRNA processing plays crucial roles in Pi homeostasis as well as developmental reprograming in response to Pi deprivation in Arabidopsis.

Foxglove aphid, Aulacorthum solani (Kaltenbach), is a Hemipteran insect that infected a wide variety of plants worldwide and caused serious yield losses in crops. The objective of this study was to identify the putative QTL for foxglove aphid resistance in wild soybean, PI 366121, (Glycine soja Sieb. and Zucc.). One hundred and forty one F2-derived F8 recombinant inbred lines developed from a cross of susceptible Williams 82 and resistant PI 366121, were used. The phenotyping of antibiosis and antixenosis was done through choice and no-choice assays with total plant damage (TPD) and primary infestation leaf damage (PLD); a genome-wide molecular linkage map was constructed with 504 single nucleotide polymorphism markers utilizing a GoldenGate assay. Using inclusive composite interval mapping analysis for foxglove aphid resistance, one major candidate QTL on chromosome 7 and 3 minor QTL regions on chromosome 3, 6 and 18 were identified. The major QTL on chromosome 7 showed both antixenosis and antibiosis resistance responses. However, the minor QTLs showed only antixenosis resistance response. The major QTL mapped to a different chromosome than the previously identified foxglove aphid resistance QTL, Raso1, from the cultivar Adams. Also, the responses to the Korea biotype foxglove aphid were different for Raso1, and the gene from PI 366121 against the Korea biotype foxglove aphid were different. Thus the foxglove aphid resistance gene from PI 366121 was determined to be an independent gene to Raso1 and designated to Raso2. This result could be useful in breeding for new foxglove aphid resistant soybean cultivars.

The adsorption characteristics of Sr and Cs ions were investigated by using PS-zeolite beads prepared by immobilizing zeolite with polysulfone (PS). The adsorption kinetics of Sr and Cs ions by PS-zeolite beads was described well by the pseudo-second-order model. The maximum adsorption capacities of Sr and Cs ions calculated from Langmuir isotherm model were 65.0 mg/g and 76.4 mg/g, respectively. In the binary system of Sr ion and Cs ion, the adsorption capacities of each ion decreased with increasing mole ratio of mixed counterpart ion, and Cs ion showed the higher hinderance than Sr ion. We found that thermodynamic properties of Sr and Cs ions on absorption reaction were spontaneous and endothermic at 293 to 323 K.

In order to recover lithium ions from aqueous solution, a novel SAN-LMO beads were prepared by immobilizing lithium manganese oxide (LMO) with styrene acrylonitrile copolymers (SAN). The optimum condition for synthesis of SAN-LMO beads was 5 g of LMO and 3 g of SAN content. The characterization of the prepared SAN-LMO beads by SEM and XRD were confirmed that LMO was immobilized in SAN-LMO beads. The removal and the distribution coefficient of lithium ions decreased with increasing lithium ion concentration and solution pH. Even when the prepared SAN-LMO beads were reused 5 times, the leakage of LMO and the damage of SAN-LMO beads was not observed.

기존의 유전자변형식물은 외래의 도입유전자를 갖고 있으며 이들로부터 기인한 단백질 또는 합성물질에 의한 의도적/비의도적 영향에 대한 안전성 논란이 사회적 이슈가 되어 왔다. 최근의 기술적 진보에 의하여 이른바 식물육종의 신기술이 발달하게 되었고 이들 기술로 만든 신규식물에 대한 안전성평가에 GMO 관련 규제의 적용 여부 문제가 대두되게 되었다. 이들 NPBTs 기술로 만든 신규식물의 특징은 SDN이나 ODM과 같이 염색체상의 정확한 위치에 짧은 염기서열의 indel(s)이나 단일염기 돌연변이를 도입하여 자연적인 돌연변이와의 구별이 거의 불가능하거나, cisgenesis와 같이 성적교잡이 가능한 종 유래의 유전자를 구조변형 없이 도입하여 근연종과 동질적인 식물을 만들거나, heterozygous 형질전환체 후대세대의 null-segregant 선발이나 epigenetic를 이용하여 도입유전자가 존재하지 않지만 목적 형질을 갖는 식물체를 만드는 장점이 있다. 또한 grafting이나 Agro-infiltration 등의 방법으로 안전성평가를 회피하거나 경감할 수 있는 가능성을 높이게 되었다. OECD를 비롯한 주요 GMO 개발국의 관련 학회에서는 SDN, ODM 및 cisgenesis 또는 intragenesis 기술로 만든 식물에 대하여 non-GM 식물과 동일한 위해성평가 규정을 적용하거나 상황에 따라 완화된 규정을 적용할 수 있다고 판단하고 있다. 현 시점에서 이들 NPBTs 기술을 이용하여 개발된 식물이 상업화된 예는 없으나 많은 국가에서 상업화를 목적으로 개발 중이며 일부에서는 안전성평가를 완료한 단계이다. 이러한 현실에서 NPBTs 기술의 개념 정립, 신규식물의 안전성평가의 방향 설정 및 현실성 있는 작물개발 방안을 마련하여야 한다. 이를 통하여 GMO에 대한 안전성 논란과 사회적 거부감을 우회하는 동시에 답보 상태에 있 는 분자육종 분야의 발전을 도모할 계기를 마련하여야 한다.

With the advent of next generation sequencers (NGS) that provide sequencing at a substantially lower cost, the development SNPs at the level of whole genomes can be done in a single laboratory. However, genome structural variation including large insertions and deletions, and chromosomal reciprocal translocations has not yet been focused due to the limitation of re-sequencing methods as genome structures rely to that of a known reference genome. For an improved detection of the structural variations after deep re-sequencing of the Glycine soja accession CS-14, we de novo assembled the whole paired-end reads (W-contigs). After the de novo assembly, the paired-end reads that did not match the reference genome of Williams 82 were retrieved and de novo assembled them (U-contigs). We then predicted structural variation candidates. For predicting the function of the structural variation candidates, we compared those structural variation candidates with SwissProt DB using BLASTX. Most of them were matched with transposable element related proteins or stress tolerance related proteins (Table 1). We designed 24 primers for all candidates and tested in CS-14 and Williams 82 for validation. As a result, the DNA polymorphism was observed between CS-14 and Williams 82 in the three primer sets, CS14IC10, CS14IC12 and CS14IC15, with the expected size of the PCR product . For further validation, we sequenced the DNA band amplified by CS14IC15, and its sequences were aligned well against the Williams 82 and CS-14 contig. Especially, IC15R-CS14 was aligned in the predicted insertion region, consequently, this sequenced region would indicate structural variation. The other primer sets did not amplified either because they were designed for an amplifying long genomic region or because of the fragmented template DNA

Korea is a origin of three basic species, P. ussuriensis, P. pyrifolia and P. fauriei. Genetic relationship among Korean pear cultivars compared with their parents were also identified that they are closely related P. pyrifolia, P. ussuriensis and/or hybrid between two species. SSRs or Microsatellites are co-dominant and typically neutral inheritance showing high degree of polymorphism, large number of alleles per locus, abundance in genomes, and suitability for automation. SSR markers were developed in apple and pear where they were used for construction of genetic linkage maps, evaluation of the genetic diversity, cultivar identification, genotype identification, and in the determination of genetic relatedness. Many apple (Malus × domestica Borkh.) SSRs would be useful for genetic mapping in European and Asian pears in previous experiments and cross-species amplification was observed between apple and pear. The objectives of this study were to develop polymorphic SSR markers in ‘Whangkeumbae’ and ‘Minibae’, which were chosen as the representative cultivars of P. pyrifolia and P. ussuriensis in each among Korean pears, from ‘Golden Delicious’ genomic sequences generated by next generation sequencing technology and to evaluate the utility of the SSR markers based on ‘Golden Delicious’ sequences. Of 51 SSR markers, 18 were polymorphic in ‘Whangkeumbae’ and ‘Minibae’. The cross-species transportability of primers designed in ‘Golden Delicious’ sequences makes SSR markers more useful, given the current high level of investment in mapping the genomes of related Rosaceae.

Bacterial spot of tomato (Solanum lycopersicum L.) is caused by at least four species of Xanthomonas with multiple physiological races. In this study, we developed a mapping population for association analysis of bacterial spot resistance. For this population, six advanced breeding lines with distinct sources of resistance were first crossed in all combinations and their F1 hybrids were intercrossed. The 1,100 segregating progeny from these crosses were evaluated in the field against T1 strains. Based on this individual evaluation, we selected 5% of the most resistant and 5% of the most susceptible progeny for evaluation as plots in two subsequent replicate field trials inoculated with T1 and T3 strains. A total of 461 markers across 12 chromosomes were used for genotyping these selections. Of these markers, an optimized subset of 384 SNPs was derived from the 7,720 SNP Infinium array developed by the Solanaceae Coordinated Agricultural Project (SolCAP). For association analysis to detect known resistance loci and additional novel loci, we used the mixed models with correction for population structure, and found that accounting for kinship appeared to be sufficient. Detection of known loci was not improved by adding a correction for structure using either a Q matrix from model-based clustering or covariate matrix from Principal Component Analysis. Both single-point and haplotype analyses identified strong associations in the region of the genome known to carry Rx-3 (chromosome 5) and Rx-4/Xv3 (chromosome 11). Additional QTL associated with resistance were detected on chromosomes 1, 3, 4, 6 and 7 for T1 resistance and chromosomes 2, 4, and 6 for T3 resistance. Haplotype analysis improved our ability to trace the origin of positive alleles. These results demonstrate that both known and novel associations can be identified using complex breeding populations that have experienced directional selection.