Poaceae, the family of grasses. One of the most successful and ubiquitous plant lineages in the world. Members of this family are known for dominating dry treeless plains, hence the name grassland. Bamboos, however, are an anomaly. Contrasting with many of their iconic relatives, bamboos represent the only major grass lineage to diversify primarily in association with forests. Their superficial tree-like appearance, built by its strong stem, also stands them out in their family of “grasses.”
禾本科，百草之族。作为植物中最成功的族群之一，它们无处不在，特别在干旱稀树的平原中尤为繁盛，于是这种群系才被叫作草原。而竹子却是一个异数，与其它禾本科不同，竹亚科似乎是唯一凭借对潮湿森林的适应而走向成功的禾本科成员。它们更是演化出了由高度木质化的茎搭建起的酷似树木的外表，在竹子矮小的亲戚中鹤立鸡群。
Not all bamboos are tall and mighty, though. As bamboos evolved from grassy ancestors (bamboos’ closest subfamily is Pooideae, containing wheat), naturally, groups that diverged early will have retained the ancestral “grassiness.” Bambusoideae, the taxonomic group bamboos belong to, has three subdivisions: the herbaceous bamboos, and two types of woody bamboos, the temperate woody bamboos and the tropical woody bamboos.
然而并不是所有的竹子都那么高大魁梧。既然木本的竹子是由草演化来的（竹子今天最近的亲戚是早熟禾亚科，包括了我们常吃的小麦），那么必定有一些分化得早的成员保留了祖先草的样子。分类学上讲，竹亚科（竹子所属的分类群）包括三种竹子：草本竹，以及两种木本竹，温带木本竹和热带木本竹。
Late Cretaceous, 70 million years ago. The group we today know as the grasses was still in its embryonic stage. Most likely in what is now South Asia, the first bamboo came to life, giving rise to the long lineage of bamboos that are to come.
七千多万年前的白垩纪，新生的禾本科还如同其它芸芸众生的类群一般，并无特别之处。而大概率在今天的南亚，也就是当时孤悬海外的印度次大陆，一株微不足道的小草的两支后代决定分道扬镳，自此竟诞生了在整个植物界中都值得称道的两大家族，禾类与竹子。
The first major event to have occurred in bamboo evolution was during the Eocene 40 million years ago. The ancestral bamboo group diverged into two lineages, the herbaceous bamboos and the woody bamboos (both temperate and tropical). Both of them continue to this day and are classified both as bamboos. Because herbaceous bamboos diverged first, it is commonly believed that they represent the ancestral appearance for bamboos.
时间来到了四千多万年前，竹子演化史上第一次大事件在始新世发生了。竹子的祖先分化出了两支后代。其中之一便是草本竹。由于它们先于其它竹子的分化，植物学家们普遍认为它们草本的身材便代表着当今木本竹早期祖先的大致形态。
Herbaceous bamboos like these used to be the most common bamboo, found throughout the whole world. However, they gradually declined, replaced by woody bamboos. Today, herbaceous bamboos can only be found in the Americas, except a single species of herbaceous bamboo living in Papua New Guinea. (Buergersiochloa bambusoides)
原先草本竹才是竹子家族中的优势种群，在全世界都有分布。然而，随着木本竹逐渐得势，草本竹也渐渐衰落，如今，除了一种分布于巴布亚新几内亚的草本竹 (Buergersiochloa bambusoides)，其余草本竹都只在美洲有分布。
While all this is happening, at Oligocene around 31 million years ago, a series of complex hybridizations (which are actually pretty common in plants) gave rise to the ancestors of modern-day temperate and tropical woody bamboos. In fact, temperate woody bamboos have a tetraploid genome, two times of the normal diploid genome, meaning it has experienced at least one mass hybridization in its evolutionary history. Some tropical woody bamboos even have hexaploid genomes, indicating at least two hybridizations.
现在让我们把注意转向三千一百万年前渐新世的木本竹。一系列的杂交事件杂交出了如今温带木本竹和热带木本竹的祖先。如今温带木本竹有着四倍体的基因组，是正常二倍体基因组的两倍，意味着在温带木本竹的演化史上至少发生过一次杂交事件。而有些热带木本竹甚至有着六倍体的基因组，即它们是经历过两次杂交后的产物。
Based on their name “temperate”, temperate woody bamboos have adapted to live in conditions of high latitudes and altitudes, so most of the bamboos we imagine covered in snow belong to temperate woody bamboos. One innovation made by the temperate bamboos is the running bamboos.
从温带木本竹的名字“温带”可以看出它们已经适应了高海拔高纬度的环境。我们脑海中所想象的被层层白雪覆盖的竹子大概率就属温带木本竹。温带木本竹有一独特的创新，即它们解锁了散生的能力。
Woody bamboos can be informally classified into two groups: clumping bamboos, or professionally called pachymorphs, and running bamboos, called leptomorphs. The ability to “run” allows one bamboo to spread rapidly and invasively, and in theory permits indeterminate extension. Not only do running bamboos possess an extraordinary ability to occupy available living space, their interconnected rhizomes also form an underground nutrition network, and the distancing between bamboos mean less competition than clumping bamboos, presenting them a significant advantage in competing with clumping bamboos. There are evidence of bamboos benefiting through deforestation by rapidly occupying those empty lands where they were previously obstructed.
所谓散生竹，是与其祖先丛生竹相对比的。散生的突破允许严格上的一个竹子向四周侵略般地生长，使形成一串盘根错节的能互相交换营养的克隆群体。而散生竹不仅能迅猛地把有限的生存空间占下，属于同一个个体的不同植株还能被分得够开而避免竞争内耗，让散生竹比起丛生竹更占优势。甚至有证据表明散生竹会利用人类的伐木迅速扩张到曾经被阻碍发展的区域。
Temperate bamboos experienced a rapid diversification around the mid-Miocene as a result of the intensification of the East Asian Monsoon, bringing a moist climate that facilitated expansion. They even crossed the Bering Land Bridge and entered North America during the Pliocene, thus extending the territory of the temperate bamboos into the Americas and bringing a seemingly alien figure to this faraway continent.
温带木本竹在中新世中叶经历了一轮多样化，而那时间段刚好是东亚季风因造山运动加剧的时候。由季风带来的潮湿气候促使了当地植物类群的辐射和扩张，乃至使温带木本竹得以跨过白令陆桥，进入北美洲安家落户，将温带竹的版图延展进了新大陆。
On the other hand, temperature fluctuations during the Oligocene-Miocene boundary caused a myriad of hybridizations (species living in different environments overlap due to those temperature fluctuations), promulgating the ancestors of the tropical woody bamboos. With the opportunity of the monsoons, they diversified from Southeast Asia both into the Americas and to Africa, where they left a lot of beautiful fossils.
在渐新世和中新世之交，频繁的温度波动促使了当时不同的竹子类群进行了进一步的杂交（因为波动的气候使得原本生长于不同环境的竹子重合了），杂交出了热带木本竹的祖先。同样是借着季风的力，它们从东南亚扩散到了美洲和非洲，并留下了许多精美的化石记录。
One of the most characteristic publicities bamboos have is the widespread trivia of “bamboos being a kind of grass.” The fact that creates this surprise is the high lignification (woodification) taking place in bamboo culm (stem) cells, which is linked to their hybrid history (still remember it?). Today, every grass has genes that code the deposit of cellulose and lignin for basic support, though at a moderate level. However, since bamboos duplicated their genomes several times due to hybridization, some of their functions are magnified as well, for example the deposit of lignin. With tetraploid or even hexaploid genomes, the deposit of lignin is multiplied to a quantity enough to make the supposed grassy stems tree-like. And this of course explains why only the polyploid bamboos happen to all be woody while the herbaceous ones are normal diploids.
如今人们经常会拿这样一个冷知识用来显摆：“竹子实际上是一类草。”而之所以人们会对此出乎意料是因为竹子茎中高度的木质化使它们和想象中的草大相径庭，这也跟它们杂交的过往有关。每种草其实多多少少都会有一些基因负责编码纤维素和木质素来保证基本的支撑。但那些木本竹的祖先在经历多倍化后，编码木质素的基因也一起翻了倍。而这就产生了量变的效果，产生的木质素也被翻了几倍。凭借着木本竹四倍体乃至六倍体的基因组，所积聚的木质素也渐渐快和树木持平了。这也完美解释了为什么多倍体的竹子都是木质的而草本的竹子都是二倍体，因为在杂交后产生的多倍体基因正是木本竹如此高大的原因。
One cool fact for bamboos is the phenomenon that all bamboos of the same species flower and breed at the same time, and after that they die simultaneously. This is because bamboo saplings are extremely shade-intolerant, and their seeds are also vulnerable against hungry herbivores. Therefore, evolution has driven them to adopt a synchronized breeding event, maximizing the success rate for the next generation of bamboos by providing a shadeless habitat and overwhelming the predators with seeds.
竹子另一有趣的点即在同一物种的竹子即使相隔千里通常会一起开花结果，并在繁殖后一同死去。这是因为竹子幼苗十分不耐荫，且掉落的竹子种子常常会成为掠食者的美餐。因此，进化使竹子同步了它们繁殖的时间，这样等它们集体死亡后，能给幼苗短暂提供一个没有任何生长阻碍的窗口，并且能靠庞大的种子数量使掠食者一时无法吃完，留下后代。
So this is the story of the bamboos. Since their establishment in the Late Cretaceous, they have persisted to this day with species numbering in the thousands and a truly global distribution. They no doubt innovated an unprecedented way of living and thrived in their own niche. And about how their future will turn out? No one knows, but we shall see.
凭借全球的分布和数千的物种量，竹子的处境似乎蒸蒸日上，且每一个竹子类群似乎都没有混得太差。从晚白垩纪自立门户起，竹子们挺着它们坚贞的形象走到了今天，在人类的文化中已经成了一种符号，一种意境。放眼异彩纷呈，争奇斗艳的植物界，竹子们无疑开辟了一类全新的生态位，至于它们的潜力与未来如何，那就让我们拭目以待吧。
For citation and a more professional version of this, see:
www.bilibili.com/read/cv21154205/
联系客服
