论文出处：
https://advances.sciencemag.org/content/7/29/eabg2286
Large-scale whole-genome resequencing unravels the domestication history of
Cannabis sativa
论文作者 https://imgur.com/J6mMzPY
Cannabis sativa has long been an important source of fiber extracted from
hemp and both medicinal and recreational drugs based on cannabinoid
compounds. Here, we investigated its poorly known domestication history using
whole-genome resequencing of 110 accessions from worldwide origins. We show
that C. sativa was first domesticated in early Neolithic times in East Asia
and that all current hemp and drug cultivars diverged from an ancestral gene
pool currently represented by feral plants and landraces in China. We
identified candidate genes associated with traits differentiating hemp and
drug cultivars, including branching pattern and cellulose/lignin
biosynthesis. We also found evidence for loss of function of genes involved
in the synthesis of the two major biochemically competing cannabinoids during
selection for increased fiber production or psychoactive properties. Our
results provide a unique global view of the domestication of C. sativa and
offer valuable genomic resources for ongoing functional and molecular
breeding research.
长期以来，大麻一直是从大麻以及基于大麻素化合物的药用和消遣性药物中提取的纤维的
重要来源。在这里，我们使用来自世界各地的 110 个种质的全基因组重测序调查了其鲜
为人知的驯化历史。我们表明C. sativa最早在东亚新石器时代早期驯化，目前所有的大
麻和药物品种都与中国目前以野生植物和地方品种为代表的祖先基因库不同。我们确定了
与区分大麻和药物品种的性状相关的候选基因，包括分支模式和纤维素/木质素生物合成
。我们还发现了在选择增加纤维产量或精神活性特性期间参与两种主要生化竞争大麻素合
成的基因功能丧失的证据。我们的结果为C. sativa的驯化提供了独特的全球视角，并为
正在进行的功能和分子育种研究提供了宝贵的基因组资源。
https://imgur.com/mZZkGgf
图表中黄色为原始大麻，绿色是纺织用麻，浅蓝色为野生迷幻药用大麻，红色是强化药性
的农业改良品种。方形是人工栽培得到的，三角形是自然杂交产生，圆形是地方特有种。
Our genome-wide analyses corroborate the existing archaeobotanical,
archaeological, and historical record [reviewed in (5, 6, 31–33)] and
provide a detailed picture of the domestication of Cannabis and its
consequences on the genetic makeup of the species. Our genomic dating
suggests that early domesticated ancestors of hemp and drug types diverged
from Basal cannabis ~12,000 years B.P. (95% confidence interval: 6458 to
15,728 years B.P.; Fig. 2B and table S3), indicating that the species had
already been domesticated by early Neolithic times. This coincides with the
dating of cord-impressed pottery from South China and Taiwan (12,000 years
B.P.), as well as pottery-associated seeds from Japan (10,000 years B.P.).
Archaeological sites with hemp-type Cannabis artifacts are consistently found
from 7500 years B.P. in China and Japan, and pollen consistent with
cultivated Cannabis was found in China more than 5000 years B.P. Only a small
number of early domesticated Cannabis strains expanded to later form hemp and
drug types ~4000 years B.P., a time when multiple fiber artifacts appear in
East Asia, and when fiber-grown Cannabis was spreading westward into Europe
and the Middle East, as shown by Bronze Age archaeological evidence.
Ritualistic and inebriant use of Cannabis has in turn been documented in
Western China from archaeological remains at least 2500 years B.P. (34, 35).
The first archaeobotanical record of C. sativa in the Indian subcontinent
dates back to ~3000 years B.P., the species likely being introduced from
China together with other crops (36, 37). In contrast with East Asia,
historical texts from India from as early as 2000 years B.P. indicate that
the species was only exploited for drug use. Over the next centuries,
drug-type Cannabis traveled to various world regions, including Africa (13th
century) and Latin America (16th century), progressively reaching North
America at the beginning of the 20th century and later, in the 1970s, from
the Indian subcontinent. Meanwhile, hemp-type cultivars were first brought to
the New World by early European colonists during the 17th century and later
replaced in North America by Chinese hemp landraces by the middle 1800s.
Consistent with this history, our model shows a gradual increase in the Ne of
hemp and drug types. On the basis of both demographic and phylogenetic
analyses, we propose that early domesticated Cannabis was first used as a
primarily multipurpose crop until ~4000 years B.P., before undergoing strong
divergent selection for increased fiber or drug production.
我们的全基因组分析证实了现有的考古植物学、考古学和历史记录
，并提供了大麻驯化及其对物种基因组成影响的详细图景。我们的基因组测年表明，
大麻和药物类型的早期驯化祖先与基础大麻约 12,000 年，表明该物种在新石器时代早期
已经被驯化。
这与来自华南和台湾的绳纹陶器（公元前 12,000 年）以及来自日本的与陶器相关的种
子（公元前 10,000 年）的年代一致。中国和日本从 7500 年 开始一直发现有大麻型
大麻文物的考古遗址，在中国发现了 5000 多年 与栽培大麻一致的花粉只有少数早期
驯化的大麻植株扩展到后来形成纺织和药物类型 ~4000 年，东亚出现多种纤维制品的
时期，以及纤维生长的大麻青铜时代的考古证据表明，它正在向西传播到欧洲和中东。
仪式和醉人使用大麻现在已经被中国西部的考古遗迹至少2500年。印度次大陆C. sativa
的第一个考古植物学记录可以追溯到大约 3000 年前，该物种可能与其他作物一起从中国
引入 。
与东亚相反，早在 2000 年 的印度历史文本表明该物种仅被用于迷幻。在接下来的几个
世纪里，迷幻类大麻前往世界各个地区，包括非洲（13 世纪）和拉丁美洲（16 世纪），
并在 20 世纪初和 1970 年代从印度次大陆逐步到达北美。与此同时，17 世纪早期的
欧洲殖民者首先将大麻品种带到了新世界，后来在 1800 年代中期在北美被中国大麻
地方品种所取代。
与这段历史一致，我们的模型显示大麻和药物类型逐渐增加。在统计学和系统发育分析
的基础上，早期驯化的大麻 直到 约 4000 年之前，它首先被用作主要的多用途作物，
然后进行极端的农业改良以增加纤维或药物产量。