馬婭·韋–哈斯 左連凱 木子

The mouth of Baishiya Karst Cave nestles near the base of a towering crag at the edge of the Tibetan Plateau. Strands of colorful prayer flags crisscross the pale face of the hollow. Within the cave’s cool confines in 1980， a local monk happened on something unexpected： a jaw with two huge teeth that， while human， was definitely not like that of humans today.

A study published in Nature reveals just how much this ancient jaw has to say. A detailed analysis of its physical features as well as proteins extracted from the fossil suggest that the mandible， dated to 160，000 years ago， comes from the enigmatic human population known as the Denisovans—a sister group to the Neanderthals previously identified from scant remains found in a single cave in Siberia’s Altai Mountains.

“I just couldn’t believe that at the moment [my colleagues] told me，” says study co-author Dongju Zhang of Lanzhou University. “I was really excited.” Zhang’s excitement is justified： the jawbone， known as the Xiahe mandible after the county where it was found， fills a yawning gap in our understanding of this mysterious ancient. While the previous Denisovan fragments come only from the eponymous Denisova Cave in Siberia， people living all across Asia and Australasia today carry Denisovan DNA in their genomes. The discovery of the Xiahe jawbone at a locale over 1，400 miles away from this Siberian cave confirms Denisovans ventured much further across the continent.

It’s thought that after the Denisovans’ ancestors split from their Neanderthal relatives at least 400，000 years ago， they headed east into Asia， while early Neanderthals spread through Eur-ope and western Asia. Modern humans first left Africa some 200，000 years ago， first as a trickle and then in waves. Eventually they encountered and interbred with Neanderthals in the Middle East. Those who trekked east into Asia likewise mated with the resident Denisovans， who left genetic fingerprints still present in Asians today.

One such Denisovan fingerprint helps modern Sherpas and Tibetans adapt to the thin air on the roof of the world. But until now， evidence of Denisovans was found only at a relatively low altitude： Denisova Cave is just 2，300 feet above sea level. The Xiahe jaw， found at roughly 10，760 feet， is the first physical evidence that Denisovans ventured up into the same altitudes as living humans carrying the genetic adaptation to low oxygen environments. And at 160，000 years old， the jawbone is four times the age of the earliest evidence of human activity in the Tibetan Plateau’s challenging climes， underscoring the remarkable resilience of our ancient relatives.

While the jaw was discovered back in the 80s， researchers only began studying it three decades later. In 2010， Lanzhou University’s Zhang， her newly minted Ph.D. in hand， turned to the strange hominin remains.

Excavations in the cave subsequently revealed large animal bones with cut marks and stone tools.

Analysis of the jaw itself turned up even more surprises. Its morphology suggests it is neither Homo erectus nor Homo sapiens， whose remains are widespread across mainland Asia. The shape of its row of teeth， for instance， was not elongated， as they are in Homo erectus. And the jaw lacks a chin—a unique trait of modern humans. Most telling of all was the sheer size of the teeth， which are similar to those from Denisova Cave bearing Denisovan DNA.

For confirmation， the researchers first attempted to extract DNA from the fossil jaw. When the analysis revealed that the ancient DNA had degraded， they turned to a more durable， if less sensitive， molecular tool： the proteins built from DNA codes. The researchers extracted proteins from both the jawbone and the tooth dentine. They then used an enzyme to cut the proteins into short strands to identify the amino acid building blocks encoded by DNA. Overall， the dentine proteins appeared much closer to the Altai Denisovans than to Neanderthals or modern humans， says Frido Welker of the University of Copenhagen， who specializes in ancient hominin proteins and led this phase of the work.

There are， however， limits to what can be said from proteins. Denisovans had a stunning amount of diversity. A study published earlier suggested that what we call Denisovans might actually be three distinct genetic lines， one of which is nearly as different from other Denisovans as they are from Neanderthals. But the similarity of proteins across groups and through generations makes it difficult to pinpoint precisely how similar the owner of the jaw is to these three Denisovan lines—or if it was from yet another sister group.

The tantalizing connection the jaw makes between what was once thought of as a low altitude group of humans and their mysterious role in modern high-altitude adaptations also remains fuzzy， explains Emilia Huerta-Sanchez， a population geneticist at Brown University and lead author of the 2014 Nature study that first identified this genetic link. “I agree with the authors that it could be that this hominin group was high altitude adapted，” she says. “But I don’t think we know for certain.”

Huerta-Sanchez explains that the genetic tweaks thought to help modern Tibetans thrive in low-oxygen environments are not part of a sequence that codes for proteins， but instead control how much of a particular protein is made. While the jaw was found where oxygen levels are low， without the DNA itself， scientists can’t be sure the jaw’s owner carried the adaptation to survive in that thin air.

While there are still many unknowns， scientists are excited about what other clues the jaw may hold for understanding human evolution in Asia. The fossil could for instance be used to identify other Denisovans from the growing pool of hominin fossils in Asia that don’t neatly fit into the known branches of our increasingly bushy family tree， says María? Martinón-Torres， who has extensively studied the Asian fossil record. For example， a three-rooted molar in the jaw is similar to that of a tooth in a previously described mandible known as Penghu 1， hinting that it， too， may be a Denisovan.

白石崖溶洞的入口靠近青藏高原邊緣高聳懸崖的底部。一串串五色經(jīng)幡交錯(cuò)于白色洞壁上。1980年，一位當(dāng)?shù)厣畟H在這個(gè)涼快的巖洞里偶然發(fā)現(xiàn)了一件意想不到的物品：帶兩顆大牙的頜骨，是人類頜骨，但肯定不是現(xiàn)代人的。

發(fā)表在《自然》雜志上的研究報(bào)告揭示了這塊古頜骨的深刻含義。對(duì)其外形特征和從化石中提取的蛋白質(zhì)的詳細(xì)分析表明，這塊16萬年前的下頜骨源于神秘的丹尼索瓦人。此前在西伯利亞阿爾泰山脈一個(gè)巖洞里發(fā)現(xiàn)的珍稀遺骨，經(jīng)鑒定確認(rèn)為尼安德特人，是丹尼索瓦人的姐妹種群。

“（同事）告訴我時(shí)，我簡(jiǎn)直不敢相信。”本研究報(bào)告共同作者、蘭州大學(xué)張東菊說，“我真的很激動(dòng)?！睆埣?dòng)得有道理：該下頜骨因其出土地夏河縣而被叫作夏河頜骨，填補(bǔ)了我們認(rèn)識(shí)這種神秘古人類的巨大空白。雖然早前只在因丹尼索瓦人而得名的西伯利亞丹尼索瓦洞里發(fā)現(xiàn)了其骨骼碎片，但當(dāng)今生活在亞洲和澳大拉西亞各地人們的基因組都攜帶著丹尼索瓦人的DNA。在距離上述西伯利亞巖洞1400英里遠(yuǎn)的地方發(fā)現(xiàn)夏河頜骨，這證實(shí)了丹尼索瓦人到過亞洲更廣闊的地方。

有人認(rèn)為，至少40萬年前，丹尼索瓦人的祖先與親緣種群尼安德特人分開后向東進(jìn)入亞洲，而早期尼安德特人分散至歐洲和西亞。大約20萬年前，現(xiàn)代人首次走出非洲，起初三三兩兩，后來成群結(jié)隊(duì)。他們最終在中東與尼安德特人相遇并生息繁衍。同樣，向東跋涉至亞洲的現(xiàn)代人與定居的丹尼索瓦人共同繁衍，從而留下仍然存在于當(dāng)代亞洲人身上的丹尼索瓦人遺傳指紋。

其中一種丹尼索瓦人遺傳指紋有助于現(xiàn)代夏爾巴人和藏族人適應(yīng)世界屋脊上的稀薄空氣。但直到現(xiàn)在，只在較低海拔處發(fā)現(xiàn)了有關(guān)丹尼索瓦人的證據(jù)：西伯利亞丹尼索瓦洞的海拔僅2300英尺。在海拔約10760英尺處發(fā)現(xiàn)的夏河頜骨，首次以實(shí)物證明丹尼索瓦人曾到達(dá)與攜帶低氧適應(yīng)基因的現(xiàn)代人同樣的海拔高度。該頜骨的主人生活在16萬年前，相比之下，目前發(fā)現(xiàn)的人類在青藏高原嚴(yán)峻氣候下活動(dòng)的最早痕跡只是在4萬年前，這進(jìn)一步說明我們的遠(yuǎn)古親戚具有非凡的適應(yīng)力。

雖然該頜骨發(fā)現(xiàn)于1980年代，但30年后才開始相關(guān)研究。2010年，蘭州大學(xué)的張東菊剛拿到博士學(xué)位，就投身研究這一陌生的古人類遺骨。

后來，在溶洞發(fā)掘出帶有切削痕跡的大塊動(dòng)物骨頭和石制工具。

對(duì)夏河頜骨本身的分析更令人吃驚。其結(jié)構(gòu)形態(tài)說明它既不屬于直立人也不屬于智人，二者的遺骸廣泛分布在亞洲大陸各地。例如，其牙齒排列的形狀不是直立人那樣的長(zhǎng)條形，并且該頜骨沒有下頦，而下頦是現(xiàn)代人獨(dú)一無二的特征。最有說服力的是其牙齒的大小，與出自丹尼索瓦洞、攜帶丹尼索瓦人DNA的那些牙齒尺寸相似。

為了確認(rèn)，研究人員首先嘗試提取化石頜骨的DNA。但分析表明古老的DNA已降解，他們轉(zhuǎn)而利用雖不敏感卻更持久的分子：由DNA代碼組成的蛋白質(zhì)。他們從頜骨和牙質(zhì)中提取蛋白質(zhì)，然后用酶將蛋白質(zhì)切割成短串，以找到由DNA編碼的氨基酸成分。哥本哈根大學(xué)的弗里多·韋爾克專門從事古人類蛋白質(zhì)的研究并領(lǐng)導(dǎo)了這一階段的工作，他認(rèn)為，總體來看，與尼安德特人或現(xiàn)代人相比，該頜骨的牙質(zhì)蛋白似乎更接近阿爾泰地區(qū)的丹尼索瓦人。

但是，蛋白質(zhì)所包含的信息是有限的。丹尼索瓦人擁有驚人的多樣性。早些時(shí)候發(fā)表的一篇論文稱，所謂的丹尼索瓦人其實(shí)可能有三個(gè)不同的遺傳譜系，其中一個(gè)譜系與其他丹尼索瓦人的差異簡(jiǎn)直跟他們與尼安德特人的差異一樣大。但種群之間和代際之間蛋白質(zhì)相似，因而很難精確說明該頜骨的主人與這三個(gè)譜系的丹尼索瓦人有多相似，抑或它是否屬于另一姐妹種群。

布朗大學(xué)的群體遺傳學(xué)家埃米莉婭·休爾塔–桑切斯解釋說，該頜骨揭示出，曾被視為生活于低海拔的人群與其對(duì)現(xiàn)代高海拔適應(yīng)基因的神秘影響之間存在令人期待的聯(lián)系，眼下尚不清晰。她帶隊(duì)首次發(fā)現(xiàn)這種遺傳聯(lián)系，并于2014年將研究成果發(fā)表在《自然》雜志上。“我和其他作者都認(rèn)為，很可能是這個(gè)古人類群體具有高海拔適應(yīng)基因?！彼f，“但我們不很確定?！?/p>

休爾塔-桑切斯解釋說，被認(rèn)為有助于現(xiàn)代藏族人在低氧環(huán)境下繁衍生息的基因變化，不存在于編碼蛋白質(zhì)的序列內(nèi)，但會(huì)控制產(chǎn)生特定蛋白質(zhì)的數(shù)量。雖然發(fā)現(xiàn)頜骨的地方氧氣稀薄，但沒有DNA，科學(xué)家無法確定，頜骨的主人是否具有在稀薄空氣里生存的適應(yīng)基因。

雖然仍有很多未解之處，但該頜骨可能對(duì)認(rèn)識(shí)亞洲人類進(jìn)化還有更多啟示，科學(xué)家對(duì)此興奮而期待。例如，廣泛研究亞洲人化石記錄的瑪麗亞·馬丁農(nóng)-托雷斯說，人類譜系不斷擴(kuò)大，越來越多的亞洲古人類化石不能與譜系的已知分支完全對(duì)應(yīng)，該化石便可用來鑒別其中的其他丹尼索瓦人。舉例來說，該頜骨上的一顆三牙根臼齒與以前發(fā)現(xiàn)的叫作“澎湖1”的頜骨上的牙齒相似，說明“澎湖1”可能也屬于丹尼索瓦人。