論文出處：
https://advances.sciencemag.org/content/7/29/eabg2286
Large-scale whole-genome resequencing unravels the domestication history of
Cannabis sativa
論文作者 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.
長期以來，大麻一直是從大麻以及基於大麻素化合物的藥用和消遣性藥物中提取的纖維的
重要來源。在這裡，我們使用來自世界各地的 110 個種質的全基因組重測序調查了其鮮
為人知的馴化歷史。我們表明C. sativa最早在東亞新石器時代早期馴化，目前所有的大
麻和藥物品種都與中國目前以野生植物和地方品種為代表的祖先基因庫不同。我們確定了
與區分大麻和藥物品種的性狀相關的候選基因，包括分支模式和纖維素/木質素生物合成
。我們還發現了在選擇增加纖維產量或精神活性特性期間參與兩種主要生化競爭大麻素合
成的基因功能喪失的證據。我們的結果為C. sativa的馴化提供了獨特的全球視角，並為
正在進行的功能和分子育種研究提供了寶貴的基因組資源。
https://imgur.com/mZZkGgf
圖表中黃色為原始大麻，綠色是紡織用麻，淺藍色為野生迷幻藥用大麻，紅色是強化藥性
的農業改良品種。方形是人工栽培得到的，三角形是自然雜交產生，圓形是地方特有種。
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.
我們的全基因組分析證實了現有的考古植物學、考古學和歷史記錄
，並提供了大麻馴化及其對物種基因組成影響的詳細圖景。我們的基因組測年表明，
大麻和藥物類型的早期馴化祖先與基礎大麻約 12,000 年，表明該物種在新石器時代早期
已經被馴化。
這與來自華南和台灣的繩紋陶器（公元前 12,000 年）以及來自日本的與陶器相關的種
子（公元前 10,000 年）的年代一致。中國和日本從 7500 年 開始一直發現有大麻型
大麻文物的考古遺址，在中國發現了 5000 多年 與栽培大麻一致的花粉只有少數早期
馴化的大麻植株擴展到後來形成紡織和藥物類型 ~4000 年，東亞出現多種纖維製品的
時期，以及纖維生長的大麻青銅時代的考古證據表明，它正在向西傳播到歐洲和中東。
儀式和醉人使用大麻現在已經被中國西部的考古遺跡至少2500年。印度次大陸C. sativa
的第一個考古植物學記錄可以追溯到大約 3000 年前，該物種可能與其他作物一起從中國
引入 。
與東亞相反，早在 2000 年 的印度歷史文本表明該物種僅被用於迷幻。在接下來的幾個
世紀裡，迷幻類大麻前往世界各個地區，包括非洲（13 世紀）和拉丁美洲（16 世紀），
並在 20 世紀初和 1970 年代從印度次大陸逐步到達北美。與此同時，17 世紀早期的
歐洲殖民者首先將大麻品種帶到了新世界，後來在 1800 年代中期在北美被中國大麻
地方品種所取代。
與這段歷史一致，我們的模型顯示大麻和藥物類型逐漸增加。在統計學和系統發育分析
的基礎上，早期馴化的大麻 直到 約 4000 年之前，它首先被用作主要的多用途作物，
然後進行極端的農業改良以增加纖維或藥物產量。