There is some important data here and details, and much for further
critical analysis. BUT: This study changes nothing. Its stated conclusions are
misleading, making false unstated assumptions and arriving at unreasonable and
unbelievable conclusions. In a way this mistake was inevitable and
obvious. The authors have concluded the the closest wild ancestors to
cultivated rice are living wild populations in the Pearl River basin.
The problem is that rice was domesticated not from living populations but from past populations almost certainly from regions where wild rice is now extinct (technically, we would say, extirpated). This study demonstrates that
big science and lots of resources do not inevitably produce answers, but that
nuanced analysis and critical thinking, and in this case some knowledge of
Chinese history, are necessary to direct analyses.
It is clear that wild rice (O. rufipogon) formerly occurred much
further north, through much of the Yangtze valley and even as far north as the
Shandong peninsula and lower Yellow River basin.
This is clearly attested from Chinese written sources of the Song Dynasty (i.e.
about 1000 years ago). Even by that period it is likely that wild rice
distribution was greatly reduced by the impact of China’s huge human population
and agricultural expansion which took place between 6000 years ago and 1000
years ago. More so than anywhere else on earth central China (from the
Yellow river to the Yangtze)
has supported massive human populations and suffered the corresponding habitat
loss. In the late Bronze Age (Zhou dynasty), they were hunting elephants on the
banks of the Yellow river (for a
wonderful book on Chinese environmental history that takes this as
representative of the broader sweep of Chinese history, see the Retreat of
the The Elephants by Mark Elvin 2004). These would certainly not be
represented in a genetic study of living elephant populations! (as blogged previously these may actually be an extinct elephant species with straight tusks)
That modern populations of Oryza rufipogon are not the direct ancestors
of japonica rice is implicit in the data in fact. The “obvious
genetic distinction between japonica and Or-IIIa (Fig. 2a)”, implies that
domestication rice and South Chinese rufipogon are in fact not really so close,
just the closest available in linving populations. The
intermediates found with Or-1 and indica are because Indian
wild rice have been less decimated by the combination climatic changes and
human impacts. Indeed this pattern is not new, but was already evidence some
years ago, especially in the study of Cheng et al 2003. (Polyphyletic origin of cultivated rice: Based on the interspersionpatterns of SINEs). —this is discussed on the basis of the more detailed Ohtsubo et al paper
or 2004 in my 2010 paper and various earlier articles in the archaeological
literature). It is nice to see a much larger dataset in the this new paper
re-affirm the results of the p-Sine study, but there is not really
anything new accept that the present authors have tries to grab a headline by
claiming a Pearl River origin for
rice. It is the populations that bridge the gap between OR-IIIa and japonica
which are crucial and these must be extinct populations of Oryza
rufipogon that were brought into cultivation in the earlier Holocene.
Geographically, this points back towards the north and the Yangtze.
The authors have found more extensive evidence that most domestication
genes were selected in japonica and then entered indica through hybridization.
Some geneticists, like the Japanese scholar Y-I Sato, or Susan McCouch at
Cornell have been discussing this for years, and evidence for this has been
mounting—you will also find discussions in the "rice consilience paper" or the "pathways to Asian civilizations" paper. It is misleading, however,
to speak of this as “introgression” which implies that pollen flow from
domesticated japonica into wild populations in India created indica. What
is missing here, and clearly absent from this study, as it was from the Molina
et al PNAS paper last year (see previous blog), is consideration of the chloroplast genome. This is
older work, but really key, because chloroplasts are not carried in pollen. The
Chloroplast (cpDNA) genome of indica and japonica are completely different.
Thus introgression by pollen flow from japonica into wild rices is a very
convoluted way to account for this hybridization as it would require
domestication genes to then persist in wild population that were then
re-domesticated. More reasobale in the model I have been promoting as the
“proto-indica”model in which wild ancestors of indica (with indica
chloroplasts) were under early cultivation or management and the were improved
by hybridization with introduced japonica. This does not require domestication
gene to somehow persist in wild population where they would be selected against
(actually I would expect such introgression to lead to the evolution of weedy
rices by "de-domestication": see this blog: ).
It also implies a role for human agency in this hybridization process. This
means that there were separate starts to cultivation (the human behaviour) for
indica and japonica even if the domestication syndrome was shared and evolved
one time.
Does genetic evidence on its own trump fossil evidence? No. Archaeological evidence, which is a fossil record of past rice and past
human activities, has once again been simply ignored! Archaeologically early
farming societies, with sedentism and villages and evidence for rice
cultivation and rice undergoing morphological changes of domestication are
found only in the Yangtze valley, as you probably well know. There is no
equivalent evidence from Guangdong/ Pearl River. In fact when rice in the Lower Yangtze
is showing morphological evolution under cultivation, i.e. between 5000 and
4000 BC, in the Pearl River and South China there are only
sparse populations of hunter-gatherer fishers, represented mainly by
coastal shell midden sites. These sites provide the earliest evidence for
ceramics in the coastal zone (more than 10,000 years later than pottery in the
Yangtze!). The first agriculture, based on rice, was introduced between 5000
and 4000 BP, although finds remain few and focused on the southern mountain
slopes and north of the Pearl River delta. By this time the Lower
Yangtze support urban sites, such as Liangzhu, support by extensive paddy field
systems and intensive cultivation of fully domesticated rice. It makes no sense
for rice domestication to be placed in the Pearl River region