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The Plant Cell 18:2855-2864 (2006) © 2006 American Society of Plant Biologists
Plant Biology Research Comes of Age in China
a Peking-Yale Joint Center of Plant Molecular Genetics and Agrobiotechnology, College of Life Sciences, Peking University, Beijing 100871, China Correspondence: xingwang.deng{at}yale.edu
A modern plant biology research enterprise is developing rapidly in mainland China. The cohort of well-trained young scientists has expanded, the prevalence of international research collaborations has risen, and the number of publications in international journals has surged over the past 10 years. In stark contrast with its limited contribution over the decades prior to 1990, China's plant biology research enterprise is more productive and internationally connected today than ever before. This trend is perhaps most apparent in China's growing representation in international journals. Here, we trace how early roots and recent reforms have laid the foundation for the development of mainland China's plant biology research enterprise and explore some of the present challenges. Overall, the number of annual publications by mainland China's plant scientists in a sample of major plant-specific and general scientific journals has increased dramatically over the last 15 years, from two publications in 1991 to 61 in 2005 (Figure 1A ). This upward trend has been particularly prominent since 2002. China's share of total Asian publications in The Plant Cell, The Plant Journal, and Plant Physiology has also grown to >20%, a dramatic increase over the previous decade (Figure 1B). However, trends in publication numbers offer only a partial glimpse of the progress that has followed broader changes in China's research environment over the past two decades. China's broad-based reform program in the late 1970s focused on revitalizing the nation's science and technology by increasing research funding and reforming institutional arrangements. With broad-based support from the government, many institutions have steadily hired new personnel and upgraded equipment and facilities. A growing number of these recruits are returning scientists trained abroad and have brought with them expertise and valuable overseas connections. These stronger international ties and China's internal experimentation with new institutional models are catalyzing fundamental changes in China's research environment.
EARLY ROOTS OF PLANT BIOLOGY IN CHINA
Modern plant biology research in China started in the 1930s, when overseas plant biologists Pei-sung Tang (trained in the U.S.) and Tsung-Le Loo (trained in Japan) returned to China and set up research and training programs in plant biology. For Tang, the program was first established at Wuhan University but influenced the later development of similar programs at other prominent universities, including National Southwest Associated University, Tsinghua University, China Agricultural University, the Chinese Academy of Sciences (CAS), Fudan University, and Peking University. His research spanned a number of areas, including plant respiration and energy production (Tang, 1981
Around the same time, Hung-chang Yin made strides in plant biology research, especially on the mechanism of chloroplast photosynthetic energy conversion, which was reported successively in two publications in Science (Yin and Sun, 1947 REFORM AND OPENING: SEEDS FOR CONTEMPORARY PLANT BIOLOGY IN CHINA China's program of reform and opening to the world in the late 1970s marked an important turning point for the sciences in China. Over the prior decade, the upheaval of the Cultural Revolution had disrupted scientific activity on a national scale. Yet after 1978, the sciences found an advocate in China's reformers. Broad-based economic reforms helped spur economic growth, which has in turn contributed to the growing funds available for research. Reforms also enabled biologists to access reagents and other materials necessary to conduct their experiments from national and international sources. Other policies focused specifically on boosting national competence in science and technology and led to funding increases, overseas training opportunities for young scientists, and infrastructure improvements.
Funding for life sciences research has grown dramatically since reforms began. The budgets of the two major funding agencies that provide the main sources of support for plant biology, the National Natural Science Foundation of China (NSFC) and Chinese Ministry of Science and Technology, have grown significantly in the past two decades (Figure 2A
; http://www.nsfc.gov.cn). As part of a long-term national strategy to raise basic and applied research capacity, life sciences research has been designated a key area for increasing future investment. As a percentage of total NSFC funding, the proportion budgeted for research in the life sciences has increased from 24 to 37% over the same period (i.e., from 19.6 million dollars in 1996 to 125.8 million dollars in 2005) (http://www.nsfc.gov.cn). On average, funding for research in the life sciences has increased by
Overseas training for young scientists has also been crucial to the revitalization of the plant sciences in China. Almost without exception, Chinese plant biologists currently engaged in notable and productive research have been trained or at least worked abroad—predominantly in North America, Western Europe, and Japan—after China launched its Open Door Policy in 1978. Many have since returned to China to take up full-time positions and establish research laboratories. Others retain their positions abroad but spend part of their time in China, where they oversee an independent research group of their own or a joint group with a local full-time faculty member. Those overseas experiences and resulting interpersonal and institutional connections have helped young scientists keep pace with the outside world, and there is an increased frequency of domestic and cross-border scientific exchange, both in terms of personnel visits and collaborations. In addition to improving training opportunities for students, closer international ties have helped to upgrade the quality of scientific publications in China. The Journal of Integrative Plant Biology (formerly Acta Botanica Sinica), which is sponsored by the Institute of Botany (CAS) and the Botanical Society of China and launched its first publication in 1952, has been published by Blackwell in the United Kingdom since 2005. The Journal of Plant Physiology and Molecular Biology, which has been sponsored by the Chinese Society for Plant Physiology and the Institute of Plant Physiology (CAS) beginning in 1964, will become an English-only journal starting in 2007 with a new name, Plant Molecular Physiology, published by Oxford University Press. Both journals used to be the major Chinese outlets for plant biology research and only published papers in Chinese. The internationalization of these journals is an encouraging development, given that publication in a journal published overseas with an overseas chief editor would have been unthinkable for many of China's scientists 10 years ago. MAJOR PLAYERS IN PLANT RESEARCH IN CHINA Institutional transformation during the reforms produced the facilities that today house most of China's plant biology research. The Shanghai Institute of Plant Physiology and Ecology (SIPPE) was formed by the integration of the former Shanghai Institute of Plant Physiology and Shanghai Institute of Entomology, CAS, in 1999 (Figure 3B ). Founded in 1944, the Shanghai Institute of Plant Physiology was one of the cradles of plant physiology research in China, and it contributed greatly to the establishment and expansion of molecular genetic research of plants and microorganisms. In particular, the institute has conducted notable research on photosynthesis and nitrogen fixation. After integration, research in SIPPE focused on functional genomics and molecular physiology of plants, microorganisms, and insects, plant–insect–microbe interactions, and molecular ecology. At the Institute of Plant Physiology and Ecology, scientists returning from overseas have provided the major driving force behind the institute's increase in research output. In the late 1990s, Xiaoya Chen (the current director), Hai Huang, and Da Luo comprised the first wave of new faculty recruits who had received advanced training abroad. The second major recruitment effort at this institute started in 2000, and a total of seven plant biology faculty members were recruited from overseas to the institute from 2000 to 2004. Most of these faculty members recruited in the last 10 years have made important contributions to their fields.
Another leading institute for plant biology research in China is the Institute of Genetics and Developmental Biology (IGDB), a CAS institute in Beijing. The IGDB was founded in 2001 by the merger of three former Institutes of CAS: The Institute of Genetics, The Institute of Developmental Biology, and The Shijiazhuang Agricultural Modernization Institute. Scientists in the IGDB work on both plant and animal models to address fundamental questions in the life sciences, focusing on areas such as genetic control of growth and development, gene expression, signal transduction, structural and functional genomics, and bioinformatics. Some researchers in the IGDB have also made significant progress on improving the water efficiency of agriculture and agronomic studies, with emphasis mainly on the improvement of crop productivity and quality. Jiayang Li, former director of the institute prior to his recent appointment as a vice president of CAS, returned to China in 1995 to launch his research program on the molecular basis of development in Arabidopsis and rice. Today, he is not only one of the leaders in the plant biology community, but also in the Chinese science community in general. A total of 11 plant biology faculty members have been recruited to this institute since 1995, and their research encompasses a wide range of areas. Peking University, the former Jing Shi Da Xue Tang (the Imperial Capital University) of the Qing Dynasty, opened in December 1898. The plant biology research program in Peking University is among the earliest in Chinese universities. Its history can be traced back to October of 1905, when the College of Agriculture was founded on the campus of the former Jing Shi Da Xue Tang. Today, 37.5% of the principal investigators at the College of Life Sciences are working in plant biology, most of whom received training abroad. Peking University has contributed to the large number of aspiring scientists trained in plant biology in China. Among them are the Peking University president Zhihong Xu and the newest returnee Hongwei Guo (who received his PhD from UCLA and postdoctoral training at the Salk Institute).
China Agricultural University (CAU) is a leading agricultural education and research institution in China, offering a wide range of subjects related to agriculture, including basic plant biology. CAU resulted from a merger of the former Beijing Agricultural University and Beijing Agricultural Engineering University in 1995. It was founded in 1949, when the College of Agriculture from Peking University, Tsinghua University, and North China University merged and formed Beijing Agricultural University. Several notable research accomplishments resulted from faculty members who have returned after receiving training abroad. For example, Dapeng Zhang's group discovered a new receptor for the plant hormone abscisic acid (ABA) (Shen et al., 2006 These four institutions (all located in Beijing or Shanghai) are perhaps the finest examples of the progress that has been made in plant biology in China, and all have realized their success by following similar paths. First, the institutes and universities share a long experience with the plant sciences. Second, they have recruited talented and well-trained young prospects from abroad. Once the recruits had arrived, they were given ample support and time to develop their research programs. In most cases, new principal investigators were usually able to begin publishing in high-profile journals after 4 to 5 years. Almost all new recruits now in these four institutions have overseas training and remain competitive and connected with the global research community. Many other institutions beyond the four mentioned above have also shown impressive progress in developing strong plant biology programs, some of which are located in Hebei, Henan, Hubei, Hunan, Zhejiang, and Guangdong provinces (Figure 3C) and will be mentioned later.
Experimentation with new institutional models has become an important strategy for identifying successful systems of research funding and support. China has recently set up a new institute for basic research in the biological sciences with a strong plant biology component, the National Institute of Biological Sciences (NIBS) at Beijing (Figure3A; www.nibs.ac.cn). Established with the full support of the Chinese leadership, NIBS serves as a new model for supporting and managing a basic research institution. The institute, under the leadership of its codirectors, Xiaodong Wang and Xing Wang Deng, operates independently without direct affiliation to any branch of government, giving it greater autonomy than traditional institutions, and is the first of its kind in China. The NIBS has its own governing body, the Board of Trustees, which comprises representatives of the related government branches and China's major scientific institutions and is responsible for institute policy and appointment of its directors. This new institute has recruited 14 full-time promising young faculty members from abroad since the first full-time faculty arrived in early 2004. At NIBS, individual research groups received full research funding for a term of 5 years, contingent upon satisfactory progress. This novel mechanism of funding support guarantees promising scientists full funding based on the quality of an applicant's past research and demonstrated capabilities. Researchers at NIBS have already shown their potential for accomplishment in their fields. A few recent examples include studies on the role of flagellin-induced defense in non-host resistance (Li et al., 2005c BREAKTHROUGHS IN RICE RESEARCH
It is hard to overestimate the importance of rice research in China, as the world's largest rice producer and consumer. China accounts for 23% of the world acreage in rice and 37% of world rice output in 2005. Domestically, rice supplies 40% of the dietary needs of the country's population of 1.3 billion (http://www.chinariceinfo.com). Longping Yuan, director of the China National Hybrid Rice Research and Development Center in Hunan Province, pioneered hybrid rice breeding and successfully developed the three-line hybrid rice breeding system in 1973. Hybrid rice now comprises
Chinese scientists also made significant contributions to sequencing and functional analysis of the rice genome. Bin Han, who was recruited as the director of the National Center of Gene Research (CAS) in 1998 and is now the vice director of the Institute of Plant Physiology and Ecology, led the sequence analysis of japonica rice chromosome 4 as part of the international rice genome sequencing effort (Feng et al., 2002
Furthermore, a number of agronomically important rice genes have been cloned and characterized by Chinese scientists. Jiayang Li's group in the IGDB have isolated and characterized the rice MONOCULM1 (MOC1) gene, which plays a critical role in controlling rice tillering and is a major determinant of rice grain yields. Their work revealed that MOC1 is specifically expressed in the axillary buds and is essential for their initiation and outgrowth (Li et al., 2003a RAPID GROWTH OF ARABIDOPSIS RESEARCH Before 1995, no Chinese scientists were doing internationally acknowledged research using the model plant Arabidopsis. Today, almost every major institute or university with a plant biology research program has multiple Arabidopsis groups, an indication that Chinese plant biologists now value both the basic and applied research in crop plants. Within the last decade, encouraging progress in Arabidopsis research has been achieved in China. Following is a brief overview of progress in key subfields.
Hormonal Signaling
Studies on ABA signaling in China also yielded important results. Although the RNA binding protein FCA has been reported as an ABA receptor involved in ABA-mediated control of flowering time, it is not the receptor for other known ABA responses (Razem et al., 2006
Light Regulation of Development
Stress Responses
Regulation by Chromatin and MicroRNA
Reproductive Development NEW DISCOVERIES IN OTHER PLANT SYSTEMS
The first article in The Plant Cell from a laboratory based in China, that of Daye Sun at Hebei Normal University, focused on the role of the heterotrimeric G protein in pollen development in lily (Ma et al., 1999
Another productive laboratory at the IGDB is headed by Yongbiao Xue, its current director, who was recruited in 1998. Yongbiao Xue's group has performed a series of interesting studies on the molecular mechanisms of self-incompatibility. In particular, his group discovered the S-locus F-box protein gene and showed that the Antirrhinum S-locus F-box protein Ah-SLF-S2 can form a SCF(AhSLF-S2) complex to target S-RNase destruction during compatible pollination but not during incompatible pollination (Qiao et al., 2004a CHALLENGES Despite these promising developments, China's modern plant biology research is in many respects still in its adolescence. In terms of research funding and output, China still lags behind many of the advanced industrialized countries, where modern plant biology research programs are far more established. There is great potential for further improvement in China, both in terms of research output and the environment for conducting plant biology research. When publication records for China and the U.S. are compared, the gap in output becomes clear. In The Plant Cell, China published 13 articles in the first 11 months of 2006, while the U.S. published 60. There is an even larger gap if the publications in the more general journals, such as Nature, Science, and Cell, are compared. Another challenge will be encouraging the contribution of female scientists, who have been underrepresented in plant sciences research in China.
As pointed out earlier, the funding for the life sciences in China is still relatively low compared with the developed countries in both the total dollar amount (Figure 2A) and the percentage of domestic GDP allocated to research and development. The overall research and development budget of China only accounts for 1.3% of the GDP, while most advanced industrialized countries with well-developed plant science research programs allocated Scientists and outside observers have pointed out that present evaluation systems for research proposals and institutional/individual performance are in need of significant modernization. The independent peer review model widely used in developed countries needs to be implemented at every level of the scientific evaluation system in China. Furthermore, the present evaluation system is typically based heavily on short-term productivity rather than long-term discovery. A more patient and balanced evaluation system on the institutional and personal productivity would likely promote risk-taking in the search for major scientific breakthroughs in the long run. FUTURE TRENDS Although challenges remain, China has made strides toward its long-term goal of modernizing its scientific enterprise. In plant biology, these efforts are perhaps most evident in the increased funding allocations, support for returning scientists, infrastructure improvements, and the strengthening of institutions and the university system. Government policies enacted during the reforms have set in motion a wave of institutional transformation and willingness to experiment that is likely to have profound effects on the way science is conducted in China by helping leaders to identify what models work best. The accomplishments of the many scientists described in this essay and others suggest that many institutes—both new and old—are growing capable of conducting research that is internationally recognized as valuable to the field.
Increased reform and openness in China have enabled Chinese scientists to take advantage of a broader trend toward the globalization of scientific research. Beyond attracting Chinese nationals, many institutions are interested in hiring the best talent regardless of nationality, and a few non-Chinese nationals have already worked in various CAS institutes. Hiring of non-Chinese nationals can be expected to increase as China's research environment becomes increasingly attractive. In contrast with the conventional perception, long-term employment of non-Chinese nationals was not uncommon throughout the Chinese history, although it was suspended for Funding for both basic and applied research in China is further expected to grow. In recent years, the funding allocated to basic research has increased, and current policies indicate that this trend will continue. 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20% per year over the last 10 years. Total research and development funding, mostly from the Ministry of Science and Technology, has increased at >20% per year over the past decade. China's research and development funding as a proportion of GDP increased from 0.6% in 1996 to 1.3% in 2005 (
