藜麦的驯化栽培与遗传育种

藜麦(Chenopodium quinoa Willd.)起源于南美洲提提喀喀湖区,是苋科(Amaranthaceae)藜属(Chenopodium)一年生作物。因其营养全面且对多种非生物逆境胁迫具有抗性,被认为是尚未被充分开发且具有高应用潜力的作物,深受育种学家的关注。近年来,随着人们对健康的关注和高品质生活的追求,对藜麦的需求量急剧增加,加之藜麦能有效缓解全球粮食安全,对其栽培与育种等研究已成为热点。为了加深对藜麦的认识和推动其产业的发展,本文结合本课题组多年来对源于安第斯山藜麦种质资源的收集、评价与利用的实践,从藜麦营养价值与应用、起源与分布、遗传研究、品种选育进展及发展趋势等方面进行了总结,以期为我国藜麦新品种(系)的培育与栽培、产业可持续发展、贫困地区人群增收及新增我国粮食生产途径等方面提供参考信息。     

藜麦基因组学与重要农艺性状位点研究进展

藜麦(Chenopodium quinoa Willd.)作为20世纪新兴的健康食物,因其营养成分全面、抗逆性强等特性备受关注,在国际上享有“营养黄金”、“素食之王”、“未来食品”的美誉。近年来随着基因组学和高通量测序技术的快速发展,藜麦高质量的全基因组序列得以完成并开展了系列关键基因功能研究。本文总结了藜麦基因组学、重要转录因子基因家族分析、遗传图谱构建和重要性状QTL定位和重要农艺和产量性状基因的研究进展。此外,针对目前藜麦育种的现状,本文还提出了藜麦育种存在的5个关键问题,并指出了未来藜麦遗传改良和育种的4个重要方向,旨在为实现未来藜麦的定向遗传改良提供参考。     

藜麦的营养成分、生物活性及加工利用

藜麦因其高营养、高生物活性而备受关注,我国许多地区早已开始了藜麦的规模化种植,但由于我国藜麦产业起步晚、研究不够深入以及加工应用的相对不足,造成藜麦产品在消费市场中仍相对空缺。为提高公众对藜麦的认识,帮助相关从业人员提高其加工水平,对藜麦的营养成分、生物活性及其加工利用现状等方面进行了概述,并简要展望了未来的研究方向和应用前景,旨在为我国藜麦加工产业的可持续发展提供新思路。     

苗菜型藜麦新品系筛选及营养品质综合评价

目的：为明确苗菜型藜麦的营养价值,进而筛选优质苗菜型藜麦新品系。方法：以30份苗菜型藜麦品系为研究材料,测定藜麦间的相关品质指标,采用SPSS 25.0进行相关性分析和系统聚类,通过模糊数学隶属函数法对各品系营养品质进行综合评价。结果：藜麦各品系营养品质差异较大。蛋白质含量的变异系数最大,为47.50%,总多酚含量的变异系数最小,为39.26%;蛋白质含量与硝态氮含量呈极显著正相关;在类间距离10处,将参试材料分为六大类,包括9个材料聚成的高蛋白高糖分类群,7个材料聚成的高蛋白低糖分类群。结论：藜麦品系JQ-01728的综合营养品质最优,可作为叶菜类蔬菜食用,也是选育苗菜型藜麦新品系的育种材料。     

基于藜麦转录组的脂肪酸生物合成途径解析

藜麦营养丰富,油脂含量高,脂肪酸组成理想,是油脂提取物的潜在资源。植物油脂主要以三酰甘油的形式储存在作物种子和果实等器官中,其合成受到环境和基因水平的调控,涉及质体、内质网和油体等多个细胞器。该文基于藜麦转录组数据,对藜麦油脂合成相关的脂肪酸生物合成途径基因进行挖掘,并对基因表达模式进行分析。结果表明:在藜麦中,与脂肪酸生物合成相关的基因序列共87条,涉及乙酰CoA羧化酶和β-酮脂酰ACP合成酶等关键酶,其中编码长链酰基辅酶A合成酶基因和β-酮脂酰ACP还原酶数目最多。通过基因表达模式分析发现,与脂肪酸生物合成相关的基因在种子表达中呈现整体上调模式,可能与种子中油脂形成和积累密切相关。对藜麦乙酰CoA羧化酶亚基编码基因进行分析发现,accD基因在不同组织间无差异表达,表明在藜麦中accD编码的β-CT亚基可能不是影响乙酰CoA羧化酶发挥作用的限制因子。藜麦KASⅡ含有保守结构域,与其他组织相比,编码基因QcFb15、QcFb45和QcFb75在种子中均存在上调表达,参与藜麦脂肪酸碳链延伸及油脂形成。对藜麦脂肪酸生物合成途径相关基因的挖掘,为藜麦油脂合成和积累的研究提供了理论基础,对高油...     

有关藜麦的一些后话

<正>藜麦,营养既均衡又全面,保健功能显著而独特;生态适应性广泛,从各方面都彰显其尊贵而不娇贵的个性。然而现状是,藜麦栽培仍然以南美洲的玻利维亚等原产区为主,远远满足不了迅速升温的藜麦消费市场需求。显然,藜麦供需的不平衡必然带来消费者不期望看到的价格的不断攀升。这个问题在20世纪末就已经凸显。随着藜麦在世界范     

“超级粮食”藜麦

<正>藜麦是一种古老的安第斯粮食作物,在蒂瓦纳科和印加文明中,藜麦在人们的饮食中占有重要地位。它对人类和动物来说都是一种营养价值很高的食物,被称为"印加稻米"。近年来,科学家们通过研究又发现了它许多隐藏的营养价值,因此藜麦也受到了全世界的关注。那么,相比其他谷物,藜麦具体有哪些较高的营养价     

盐胁迫对藜麦种子萌发的影响研究

再生水资源可浇灌农田,但水中含有的阴离子可使土壤产生盐胁迫。为研究盐胁迫对藜麦(Chenopodium quinoa)种子萌发特性及胚根、胚芽生长的影响,该研究以6个藜麦品种(红藜麦、国红藜麦、台红藜麦、台紫红藜麦、黄藜麦、台黄红藜麦)为材料,分别以NaCl、Na₂SO₄、NaHCO₃和对照(CK)处理6个藜麦品种种子,测定其发芽率、胚根、胚芽抑制率等指标,运用均方差决策法对不同藜麦品种耐盐性进行综合评价,初步筛选出不同盐胁迫下耐盐性较强的品种。结果表明：(1)三种盐胁迫中,Na₂SO₄对种子萌发指标抑制作用最明显,6个藜麦品种的发芽率均相对较低,一直保持在5%以下,除黄藜麦、台黄红藜麦,其余4个品种的活力指数和生长速率均为0,除黄藜麦外,Na₂SO₄ 对其余5个藜麦品种的胚根、胚芽抑制率均达到100%;NaCl对种子萌发和生长的抑制作用较小,甚至可促进胚根和胚芽生长,国红藜麦和台黄红藜麦的生长速率在NaCl处理下始终高于对照,在9 ...     

基于荧光原位杂交的藜属植物核型分析

为精确地识别藜属植物染色体组的核型特征,该文研究了4种来自青海高原的野生藜属植物(灰绿藜、藜、菊叶香藜及杂配藜)和1种从美国引进的栽培藜麦品种PI614932-HX(3)基于染色体荧光原位杂交(rDNA FISH)的核型。利用5S rDNA和45S rDNA对5种藜属植物有丝分裂中期的染色体进行FISH研究。藜属植物的核型分析结果表明:(1)藜属植物中存在二倍体(2n=2x=18)和四倍体(2n=4x=36)两种倍性,藜麦和灰绿藜为四倍体,其余3种为二倍体。(2)藜麦、灰绿藜、藜、菊叶香藜及杂配藜的核型公式分别为2n=4x=36=34m(2AST)+2sm,2n=4x=36=32m(4AST)+4sm,2n=2x=18=16m(4AST)+2sm,2n=2x=18=18m及2n=2x=18=16m+2sm。(3)染色体由大部分的中部着丝粒染色体(m)和少部分近中部着丝粒染色体(sm)组成。(4)核型类型除了菊叶香藜为1B以外,其余均属于2B类型。(5)在藜麦、灰绿藜及藜中具有分布位置不同、数量不等的双随体。5S rDNA、45S rDNA FISH结果表明:(1)藜麦和灰绿藜的染色体上存在2对5S rDNA位点和1对45S rDNA位点,藜、杂配藜的染色体上存在1对5S rDNA位点和1对45S rDNA位点,菊叶香藜的染色体上只存在1对5S rDNA位点。(2)5S rDNA和45S rDNA位点均位于染色体的短臂上。该研究首次获得了藜属植物基于5S rDNA和45S rDNA荧光原位杂交核型,为藜属植物亲缘关系研究和细胞生物学研究提供了分子细胞遗传学依据。     

藜麦宝藏如何藏

<正>藜麦是异源四倍体植物,分布于美洲热带和亚热带地区。在对干旱、盐碱地、高原和其他边缘环境漫长的适应过程中,形成了丰富的种质多样性,众多藜麦野生品种资源习性和性状参数的变化,也成就了藜麦的千姿百态。藜麦的生长习性可以分为4种:不分枝,只有单一主穗;分枝顶穗长至主穗下部的三分之一     

Assessment of genetic diversity patterns in Chilean quinoa (Chenopodium quinoa Willd.) germplasm using multiplex fluorescent microsatellite markers

Quinoa (Chenopodium quinoa Willd.) is a staple seed crop in the Andean region of South America. Improving quinoa productivity is a primary food-security issue for this region, and has been part of the impetus for the establishment of several new quinoa breeding programs throughout the Andean region. Chilean quinoa has been characterized as morphologically diverse and bifurcated into coastal and highland ecotypes. The success of emerging breeding programs will rely heavily on the development of core germplasm collections and germplasm evaluation—especially of the coastal quinoa ecotypes that are often neglected in traditional breeding programs. Thus, the objective of this study was to characterize and quantify the genetic diversity within 28 Altiplano and 31 coastal Chilean accessions of quinoa using microsatellite markers. To facilitate the analysis, we also report the development of seven sets of fluorescent multiplexed microsatellite PCR reactions that result in genetic information for 20 highly polymorphic microsatellite loci. A total of 150 alleles were detected among the quinoa accession, ranging from 2 to 20 alleles per locus and an average 7.5 allele/locus. Both cluster (UPGMA) and principal component analyses separated the accessions into two discrete groups. The first group contained quinoa accessions from the north (Andean highlands) and the second group consisted of accessions from the south (lowland or coastal). Three accessions from Europe were classified into the southern quinoa group. The data obtained in the diversity analyses highlights the relationships within and among northern and southern Chilean quinoa accessions and provides the quinoa scientific community with a new set of easy to use and highly informative genetic markers.     

Molecular and cytological characterization of ribosomal RNA genes in Chenopodium quinoa and Chenopodium berlandieri.

The nucleolus organizer region (NOR) and 5S ribosomal RNA (rRNA) genes are valuable as chromosome landmarks and in evolutionary studies. The NOR intergenic spacers (IGS) and 5S rRNA nontranscribed spacers (NTS) were PCR-amplified and sequenced from 5 cultivars of the Andean grain crop quinoa (Chenopodium quinoa Willd., 2n = 4x = 36) and a related wild ancestor (C. berlandieri Moq. subsp. zschackei (Murr) A. Zobel, 2n = 4x = 36). Length heterogeneity observed in the IGS resulted from copy number difference in subrepeat elements, small re arrangements, and species-specific indels, though the general sequence composition of the 2 species was highly similar. Fifteen of the 41 sequence polymorphisms identified among the C. quinoa lines were synapomorphic and clearly differentiated the highland and lowland ecotypes. Analysis of the NTS sequences revealed 2 basic NTS sequence classes that likely originated from the 2 allopolyploid subgenomes of C. quinoa. Fluorescence in situ hybridization (FISH) analysis showed that C. quinoa possesses an interstitial and a terminal pair of 5S rRNA loci and only 1 pair of NOR, suggesting a reduction in the number of rRNA loci during the evolution of this species. C. berlandieri exhibited variation in both NOR and 5S rRNA loci without changes in ploidy.     

Breeding quinoa (Chenopodium quinoa Willd.): potential and perspectives

Quinoa (Chenopodium quinoa Willd.) originated in the Andean region of South America; this species is associated with exceptional grain nutritional quality and is highly valued for its ability to tolerate abiotic stresses. However, its introduction outside the Andes has yet to take off on a large scale. In the Andes, quinoa has until recently been marginally grown by small-scale Andean farmers, leading to minor interest in the crop from urban consumers and the industry. Quinoa breeding programs were not initiated until the 1960s in the Andes, and elsewhere from the 1970s onwards. New molecular tools available for the existing quinoa breeding programs, which are critically examined in this review, will enable us to tackle the limitations of allotetraploidy and genetic specificities. The recent progress, together with the declaration of “The International Year of the Quinoa” by the Food and Agriculture Organization of the United Nations, anticipates a bright future for this ancient species.     

Phenological growth stages of quinoa (Chenopodium quinoa) based on the BBCH scale

Quinoa (Chenopodium quinoa) is a pseudocereal native from the Andean region of South America that has increased in importance worldwide. Quinoa is now considered an alternative to traditional crops in a climate change scenario, considering its ability to adapt to marginal soils, droughts and frosts. Despite the interesting agronomic and nutritional features of this crop, research into quinoa is characterised by individual attempts to define its phenological stages without an international consensus. A unique criterion to quantify the phenology of quinoa could become a useful tool for researchers and plant breeders in future work by standardising this information for international cooperation. In this article, a proposed scale of the phenological growth stages of quinoa based on the BBCH coding system (Biologische Bundesanstalt Bundessortenamt und CHemische Industrie) was developed. Growth stages were described utilising the decimal code of the BBCH system, and figures were included for the most representative stages.     

A recessive allele inhibiting saponin synthesis in two lines of Bolivian quinoa (Chenopodium quinoa Willd.)

Quinoa cultivars currently grown in North America and Europe require removal of bitter-tasting saponins from the grain prior to human consumption. This need for postharvest processing is a barrier to expanding production of the crop outside its Andean area of origin. Grain saponin content in quinoa shows continuous variation and is considered to be a quantitative trait. However, segregation for the presence or absence of grain saponin in F2 generations derived from crosses between high- and low-saponin parents indicates a major gene effect, with plants homozygous for a recessive allele spl having no detectable grain saponin. Variation in saponin levels among F2 plants with detectable grain saponin was consistent with polygenic inheritance. It appears that grain saponin level in quinoa is both qualitatively and quantitatively controlled, with saponin production requiring at least one dominant allele at the Sp locus and the amount of grain saponin being determined by an unknown number of additional quantitative loci. Introgression of sp1 into day-neutral lines will facilitate the development of short-season "sweet" quinoa cultivars which do not require postharvest processing to remove grain saponin.     

Quinoa – a Model Crop for Understanding Salt-tolerance Mechanisms in Halophytes

Quinoa (Chenopodium quinoa Willd.) is an ancient Andean crop that produces edible seeds and leaves. Quinoa's tolerance to salinity and other types of abiotic stresses provides it with high potential in a world where scarcity of water and increased soil salinization are important causes of crop failures. Due to its traditionally broad cultivation area (from Colombia to southern Chile), there is a wide range of quinoa cultivars adapted to specific conditions displaying a broad genetic variability in stress tolerance. In addition, being practically unique as a halophytic seed-producing crop with amazing nutritional properties, it is ideal as a model species for investigating morphological, cellular, physiological, and bio-molecular mechanisms of salinity tolerance. This review summarizes current knowledge of genotype-dependent variability in salinity responses and adaptive salt-tolerance mechanisms in quinoa. These include anatomical features and physiological aspects, such as osmotic adjustment through accumulation of ions, osmoprotectants, and sodium loading, transport, and storage, including the activity and gene expression of plasma and vacuolar membrane transporters. Finally, current knowledge regarding the effect of salinity on the nutritional properties of quinoa is discussed.     

Simple sequence repeat marker development and genetic mapping in quinoa (Chenopodium quinoa Willd.)

Quinoa is a regionally important grain crop in the Andean region of South America. Recently quinoa has gained international attention for its high nutritional value and tolerances of extreme abiotic stresses. DNA markers and linkage maps are important tools for germplasm conservation and crop improvement programmes. Here we report the development of 216 new polymorphic SSR (simple sequence repeats) markers from libraries enriched for GA, CAA and AAT repeats, as well as 6 SSR markers developed from bacterial artificial chromosome-end sequences (BES-SSRs). Heterozygosity (H) values of the SSR markers ranges from 0.12 to 0.90, with an average value of 0.57. A linkage map was constructed for a newly developed recombinant inbred lines (RIL) population using these SSR markers. Additional markers, including amplified fragment length polymorphisms (AFLPs), two 11S seed storage protein loci, and the nucleolar organizing region (NOR), were also placed on the linkage map. The linkage map presented here is the first SSR-based map in quinoa and contains 275 markers, including 200 SSR. The map consists of 38 linkage groups (LGs) covering 913 cM. Segregation distortion was observed in the mapping population for several marker loci, indicating possible chromosomal regions associated with selection or gametophytic lethality. As this map is based primarily on simple and easily-transferable SSR markers, it will be particularly valuable for research in laboratories in Andean regions of South America.     

Genome size variation in Chenopodium quinoa (Chenopodiaceae)

The extent and significance of intraspecific genome size variation were analysed in quinoa (Chenopodium quinoa Willd.), a pseudocereal important for human consumption in the Andean region of South America. Flow cytometry, with propidium iodide as the DNA stain, was used to estimate the genome size of 20 quinoa accessions from Ecuador, Peru, Bolivia, Argentina, Chile and the USA. Limited genome size variation was found among the analysed accessions. The differences between the accessions were statistically significant but the maximum inter-accession difference between the populations with the largest and the smallest genome reached only 5.9%. The largest genome was found in population C4 from Chile (mean 3.077 pg/2C) and the smallest in the Peruvian population P2 (mean 2.905 pg/2C). The variation was not correlated with collection site; however, the quinoa accessions analysed in this study belonged to three distinct geographical groups: northern highland, southern highland and lowland.