© The Authors, 2026, Published by the Universidad del Zulia*Corresponding author: lichao888@jlnku.edu.cn; hb@jlnku.edu.cn
Keywords:
Japan
Rice
Transgenic technology
Agrobacterium-mediated transformation
Genetic transformation system
Review
Contributions and application advances of japan in the development of rice genetic
transformation technology systems
Aportes y avances en la aplicación de japón en el desarrollo de sistemas de tecnología de
transformación genética del arroz
Contribuições e avanços na aplicação do japão no desenvolvimento de sistemas de tecnologia de
transformação genética do arroz
Taiye Lai
Chao Li
Bing He*
Xiaohang Wang
Hong Lang
Shuai Wang
Rev. Fac. Agron. (LUZ). 2026, 43(3): e264336
ISSN 2477-9407
DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v43.n3.IV
Crop production
Associate editor: Dr. Jorge Vilchez-Perozo
University of Zulia, Faculty of Agronomy
Bolivarian Republic of Venezuela
College of Agriculture, Jilin Agricultural Science and
Technology University, Jilin City, Jilin Province, China,
132101.
Received: 03-04-2026
Accepted: 12-06-2026
Published: 28-06-2026
Abstract
Based on a systematic literature retrieval from multiple
mainstream databases including Web of Science, Google Scholar
and J-STAGE, this paper collected and sorted relevant research
literature published from 1960 to 2024, and also supplemented
information through exchanges with Japanese rice biotechnology
researchers. Japan has made foundational contributions to rice
genetic transformation technology, establishing key systems
such as protoplast regeneration and Agrobacterium-mediated
transformation. These advances have enabled the development of
transgenic rice with improved stress resistance, enhanced quality,
and nutritional fortication. Despite slow commercialization
domestically, Japan’s technological platforms have become
essential tools for global functional genomics and molecular
breeding in rice. The integration of these systems with gene editing
promises innovative solutions for future food security challenges.
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Rev. Fac. Agron. (LUZ). 2026, 43(3): e264300 July-September ISSN 2477-9407.
2-5 |
Resumen
Mediante la búsqueda sistemática de literatura en las principales
bases de datos como Web of Science, Google Scholar y J-STAGE, se
recopilaron y clasicaron los documentos de investigación publicados
entre 1960 y 2024; además, se complementó la información a
través de intercambios con investigadores japoneses especializados
en biotecnología del arroz. Japón ha realizado contribuciones
fundamentales a la tecnología de transformación genética del arroz,
estableciendo sistemas clave como la regeneración de protoplastos
y la transformación mediada por Agrobacterium. Estos avances han
permitido el desarrollo de arroz transgénico con mayor resistencia al
estrés, calidad mejorada y forticación nutricional. A pesar de una
lenta comercialización a nivel nacional, las plataformas tecnológicas
de Japón se han convertido en herramientas esenciales para la
genómica funcional global y el mejoramiento molecular en arroz.
La integración de estos sistemas con la edición genética promete
soluciones innovadoras para los futuros desafíos de seguridad
alimentaria.
Palabras clave: Japón, arroz, tecnología transgénica, transformación
mediada por Agrobacterium, sistema de transformación genética.
Resumo
Com base na busca sistemática de literatura em diversas bases
de dados consolidadas, incluindo Web of Science, Google Scholar
e J-STAGE, foram reunidos e organizados os trabalhos de pesquisa
publicados entre 1960 e 2024, e as informações foram ainda
enriquecidas por meio de comunicações com pesquisadores japoneses
da área de biotecnologia do arroz. O Japão realizou contribuições
fundamentais para a tecnologia de transformação genética do arroz,
estabelecendo sistemas-chave como a regeneração de protoplastos e a
transformação mediada por Agrobacterium. Esses avanços permitiram
o desenvolvimento de arroz transgénico com maior resistência a
estresses, qualidade aprimorada e forticação nutricional. Apesar de
uma lenta comercialização doméstica, as plataformas tecnológicas do
Japão tornaram-se ferramentas essenciais para a genómica funcional
global e o melhoramento molecular do arroz. A integração desses
sistemas com a edição genética promete soluções inovadoras para os
futuros desaos de segurança alimentar.
Palavras-chave: Japão, arroz, tecnologia transgénica, transformação
mediada por Agrobacterium, sistema de transformação genética.
Introduction
Against the backdrop of population growth, limited arable land, and
climate change, conventional rice breeding methods face bottlenecks
such as long cycles and limited eciency in improving complex
agronomic traits (Ahmar et al., 2020). In this context, transgenic
technology has become an important means to accelerate crop
genetic improvement and achieve precise trait design. Internationally,
transgenic crops such as maize and soybean are widely used, while
transgenic research on rice, serving as both a model crop and a key
food grain, holds both scientic and strategic signicance.
As a leader in agricultural science and technology, Japan
possesses a profound accumulation in both fundamental rice research
and breeding practices (Li et al., 2023). Since the mid-20th century,
Japanese scientists have made pioneering breakthroughs in key
rice technologies like tissue culture and genetic transformation.
These advances have laid a methodological foundation for domestic
biotechnology research and provided essential technological
platforms for global rice functional genomics and molecular breeding
(Nakagahra et al., 1997).
This article reviews Japan’s key contributions and the evolution
of its transgenic rice research, focusing on the established genetic
transformation systems and their derived applications. This review
is intended to provide a reference for a deeper understanding of the
development path of rice transgenic technology and its role in food
security.
Methods
This review is based on a systematic retrieval and analysis of peer-
reviewed journal articles published from the 1960s to 2024, sourced
from databases such as Web of Science, Scopus, PubMed, Google
Scholar, and J-STAGE. To supplement and corroborate the literature
information, the writing of this review also referred to indirect
communication with several Japanese scientists long engaged in rice
biotechnology research and an analysis of viewpoints from important
review authors in related elds, aiming for a more comprehensive
understanding of the historical context and research challenges in
technological development.
Discussion
Callus Induction and Plant Regeneration
The theoretical foundation of plant tissue culture comes from the
concept of “totipotency,” rst proposed by Gottlieb Haberlandt in
the early 20th century. Building on this concept, Hiroshi Niizeki and
Kiyoshi Oono achieved the rst successful rice anther culture in 1968,
producing haploid plants and marking the beginning of rice tissue
culture research (Niizeki & Oono, 1968). This milestone subsequently
enabled the development of doubled haploid (DH) production through
anther culture. In the 1970s, rapid optimization of anther culture
conditions ensued, including cold pretreatment of materials before/
after heading, control of donor plant physiological status, adjustment
of sugar concentration and hormone ratios (e.g., 2,4-D, cytokinins),
and selection of culture medium systems. Genotype dependency was
claried (Japanese japonica rice generally responds more readily
than indica rice) (Komamine, 2003). Anther culture entered a trial
stage in breeding for Japanese japonica materials, forming an early
methodological basis for rapid DH line xation. During 1978-1980,
Japanese research groups systematically established an embryogenic
callus induction and plant regeneration system using immature
embryos (specically the scutellum) as explants, promoting the
routine regeneration of plants from somatic starting materials (Radi
& Maeda, 1987). Ozawa and Komamine (1989) conducted early
and systematic histological and physiological studies on somatic
embryogenesis in rice, providing theoretical and technical support for
high-frequency regeneration.
Concurrently, explorations into protoplast culture and regeneration
were undertaken, systematically evaluating the manifestation and
utilization potential of “somaclonal variation” in rice (e.g., obtaining
resistance or quality trait variations) (Sun et al., 1991; Zong-Xiu et
al., 1983). In the 1990s, stable and ecient regeneration systems
based on embryogenic calli induced from mature/immature seeds
were established, paving the way for gene function research.
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Lai et al. Rev. Fac. Agron. (LUZ). 2026, 43(3): e264300
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Plant regeneration from protoplasts and its role in rice genetic
transformation
The rst successful regeneration of whole plants from rice
protoplasts marked a signicant breakthrough in protoplast culture
for cereal crops. Fujimura et al. (1985) were the rst to regenerate
complete plants from protoplasts isolated from rice suspension cells,
demonstrating that establishing embryogenic suspension cell lines
with strong regenerative capacity was key to achieving protoplast
regeneration. Subsequently, Yamada et al. (1986) and Toriyama
et al. (1988) also achieved plant regeneration from protoplasts in
several Japanese rice cultivars, further conrming the applicability
of this technique across dierent genotypes. To improve regeneration
eciency, Kyozuka et al. (1987) developed a novel nurse culture
method. This method combined agarose bead culture with actively
growing nurse cells, achieving a colony formation frequency of up
to 10 % from protoplasts and a plant regeneration frequency of 17
%-50 %, regenerating numerous transplantable plants via somatic
embryogenesis. The establishment of this technique transformed
rice protoplast culture from a laboratory exploration into a stable,
reproducible operational system, laying a solid foundation for
subsequent genetic manipulation.
With the maturation of the protoplast regeneration system,
Japanese researchers quickly applied it to the direct introduction
of foreign genes, pioneering rice transgenic technology. Uchimiya
et al. (1986) rst used the polyethylene glycol (PEG) method to
introduce a plasmid carrying a kanamycin resistance gene into rice
protoplasts and obtained transgenic calli via antibiotic selection, with
Southern blotting conrming foreign gene integration. Shortly after,
Toriyama et al. (1988) successfully regenerated fertile transgenic rice
plants from protoplasts into which a kanamycin resistance gene had
been introduced via electroporation, accomplishing the full process
from gene introduction to whole plant regeneration. Thereafter,
Shimamoto et al. (1989) further optimized electroporation conditions,
co-introducing a hygromycin phosphotransferase (hph) gene and a
β-glucuronidase (GUS) reporter gene into protoplasts, obtaining
transgenic rice stably expressing the foreign genes, and conrming
stable inheritance through progeny analysis. Collectively, these
studies demonstrated that obtaining transgenic plants via direct gene
introduction into protoplasts had become a stable technique for rice
genetic transformation.
Rice genetic transformation using Agrobacterium
Although transgenic technology had been established in rice
via direct gene delivery methods into protoplasts (e.g., PEG,
electroporation), this approach had signicant limitations: many rice
cultivars (especially indica) were dicult to culture as protoplasts,
regeneration capacity varied greatly among cultivars, and regenerated
plants often exhibited high deformity rates and diculty in obtaining
normal fertile plants; furthermore, protoplast culture technology was
complex, time-consuming, and required highly skilled operation. The
gene gun method (particle bombardment) partly circumvented the
diculties of protoplast culture but still suered from issues such
as high copy number integration and frequent DNA rearrangements.
Concurrently, due to dierences in monocot cell wall structure and
phenolic compound secretion, the natural infection capability of
Agrobacterium tumefaciens towards them was very weak, leading to
the long-held belief that Agrobacterium-mediated gene transfer was
dicult to achieve in rice. In 1994, Japanese scientists rst broke
through this bottleneck. Using embryogenic calli (primarily derived
from immature embryo scutella) as recipients, they successfully
obtained a large number of morphologically normal, fertile transgenic
rice plants by optimizing Agrobacterium strains, co-culture conditions,
and selection strategies, achieving transformation eciencies
comparable to those in dicot plants (Hiei et al., 1994). This study rst
proved that Agrobacterium could eciently transfer T-DNA into rice
cells with stable integration, expression, and inheritance, establishing
a milestone technological foundation for rice genetic transformation.
Building upon the breakthrough by Hiei et al. (2008), Japanese
teams further promoted vector and transformation system
optimization. Komari et al. (2006) developed the super-binary vector,
which incorporated additional virulence genes (e.g., virB, virG from
pTiBo542) on the basis of standard binary vectors, signicantly
enhancing T-DNA transfer eciency, particularly showing high
transformation eciency for dicult-to-transform japonica cultivars
(e.g., ‘Koshihikari’) and some indica materials. Subsequently,
Hiei and Komari (2008) systematically summarized an ecient
transformation protocol using calli induced from immature or
mature embryos as recipients. This protocol, through heat shock and
centrifugation pretreatment, addition of suitable phenolic inducers
(e.g., acetosyringone), and optimized selection procedures, achieved
high-eciency transformation (50 %-90 %) for various genotypes,
including japonica and indica (e.g., ‘Kasalath’). This standardized
protocol became a core technological platform for global rice
functional genomics research and genetic improvement.
Agrobacterium-mediated transformation became the mainstream
method, replacing direct delivery methods (Wakita et al., 1998),
due to its distinct advantages: Direct delivery methods (e.g., gene
gun) often lead to complex integration of foreign genes as multiple
copies, rearrangements, or fragments, prone to causing gene silencing
or unstable expression. In contrast, Agrobacterium-mediated
transformation typically results in low-copy (1-2), precise T-DNA
integration, favoring stable transgene expression and inheritance.
The Agrobacterium method does not require expensive specialized
equipment (e.g., gene gun) and can leverage extensive experience in
vector construction and strain manipulation accumulated in dicots,
presenting a relatively lower technical barrier. Using embryogenic
calli as recipients avoids the genotype dependency and regeneration
diculties associated with protoplast culture, making it particularly
suitable for indica cultivars that are dicult to regenerate from
protoplasts.
Diversied systems for rice genetic transformation
Following the establishment of the Agrobacterium-mediated
transformation system pioneered by Japanese teams as the core
platform for global rice functional genomics and molecular breeding,
its technical procedures gradually became standardized and scaled up
in the 2010s. This ecient and stable regeneration system, particularly
protocols based on easily regenerable model japonica cultivars
like ‘Nipponbare’ and ‘Kitaake’, provided the indispensable gene
delivery and plant regeneration steps for the ecient implementation
of CRISPR/Cas gene editing technologies in rice (Sukegawa et al.,
2023). Japanese researchers continuously optimized culture medium
hormone ratios, light/temperature conditions, and osmotic regulation,
eectively shortening culture cycles, reducing somaclonal variation,
and improving the uniformity of regenerated plants (Ozawa, 2012). To
overcome the bottleneck of dicult regeneration in some indica and
local cultivars, emerging strategies such as introducing developmental
regulator genes (e.g., BBM, WUS) or their transient expression
systems were employed to directly enhance the regenerative capacity
of recipient cells, further expanding the genotype applicability of the
Agrobacterium method (Chen et al., 2022).
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Rev. Fac. Agron. (LUZ). 2026, 43(3): e264300 July-September ISSN 2477-9407.
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Despite the widespread adoption of Agrobacterium-mediated
transformation, research into simpler and less expensive
transformation methods is still ongoing. Among these, silicon carbide
whisker (SCW)-mediated transformation oers a unique pathway.
SCWs are ne needle-like crystals (Karasek et al., 1989). The
principle involves vortex-mixing plasmid DNA, SCWs, and plant
cells (e.g., embryogenic tissues), where the whiskers create micro-
wounds on cell surfaces, facilitating direct DNA uptake (Asad &
Arshad, 2011). This method has been successfully applied in crops
like maize. Notably, Japanese research teams have successfully
achieved SCW-mediated rice genetic transformation using the same
mature embryo-derived scutellum tissues as recipients commonly
used in gene gun methods (Matsushita et al., 1999). The greatest
advantage of this method is that it does not require expensive
specialized gene delivery equipment (e.g., gene gun) nor complex
protoplast or callus culture systems; its operation is extremely simple.
It is regarded as one of the most promising simplied transformation
schemes, particularly suitable for application in developing countries
with limited laboratory resources.
Application directions and representative genes in transgenic
rice
In the early 1990s, rice transgenic technology began transitioning
from methodological exploration to practical application, and for the
rst time, exogenous genes with clear practical value were introduced
into rice, marking the entry of rice genetic engineering breeding into
a substantial stage. Subsequently, research utilizing this technology
for rice improvement expanded extensively across multiple
directions, evolving from initial focus on enhancing resistance to
comprehensively improving quality, strengthening environmental
adaptability, and endowing rice with novel functions.
Resistance improvement targeting major production
challenges
Early applications focused on addressing key pest, disease, and
weed problems. Researchers successfully introduced the rice stripe
virus coat protein gene into rice, developing resistant materials, with
some lines completing environmental safety assessments and entering
general farmland cultivation (Satoh et al., 2010). Concurrently,
the development of insect-resistant rice by introducing Bacillus
thuringiensis
(Bt) endotoxin genes was also achieved (Fujimoto et
al., 1993). Regarding herbicide resistance, research was not limited
to single herbicides but explored introducing genes with broad-
spectrum detoxication potential to achieve wider eld adaptability
(Kawahigashi et al., 2003).
Quality optimization for consumer demand
To directly improve rice grain quality, genetic engineering was
applied to regulate key components. Using antisense technology
to suppress the expression of the rice Waxy gene aimed to reduce
amylose content, breeding high-gluten rice suitable for specic
dietary needs (Isshiki et al., 1998). Similarly, suppressing storage
protein glutelin genes via antisense technology could reduce grain
protein content, aiming to breed raw material rice more suitable for
brewing (Takaiwa et al., 1999; Washida et al., 1999). Addressing
food safety, transgenic rice with suppressed expression of the major
rice allergen 16kD albumin via antisense genes was developed, and
related low-allergen rice has entered the practical application testing
stage (Nakase et al., 1997).
Integrated strategies for enhanced abiotic stress tolerance
To cope with complex environmental stresses, research shifted
towards traits involving intricate physiological pathways. For
example, genetically engineering enhanced synthesis of glycine
betaine not only signicantly improved rice salt tolerance but also
synergistically enhanced adaptation to multiple stresses including
drought, low temperature, and high temperature (Hayashi & Murata,
1998). Furthermore, attempts were made to improve temperature
adaptation in rice by altering membrane system-related properties
such as fatty acid composition (Matsumura et al., 2002; Takeuchi et
al., 2001).
Yield and physiological function optimization for the future
To further explore yield potential and address global climate
change, research delved into core physiological processes like
photosynthesis. By introducing various genes related to plant defense
responses, their eects on enhancing pest and disease resistance
were evaluated (Sharoni et al., 2011; Yamaguchi et al., 2009).
Simultaneously, genetic modication research aiming to increase
yield and optimize photosynthetic capacity to adapt to global warming
was also conducted (Matsuoka et al., 2001; Suzuki et al., 2000).
Innovative exploration for novel functions in rice
Going beyond the scope of traditional breeding, genetic
engineering was employed to endow rice with entirely new production
functions. Examples include breeding rice capable of producing
soybean protein (Katsube et al., 1999), iron-fortied rice with high
iron content (Masuda et al., 2013), and functional rice containing
specic immunity-enhancing substances (Suzuki et al., 2003).
In summary, genetic engineering technology has enabled a leap
in rice breeding objectives—from single resistance to comprehensive
superior traits, from environmental adaptation to the creation of novel
functions. Rice combining high environmental resilience, strong pest/
disease resistance, and enhanced nutritional value holds promise for
achieving stable high yields under adverse conditions, oering more
possibilities for addressing future food security challenges. As more
novel genes with application potential are isolated and characterized,
and evaluated using mature transformation systems, the value of
genetic engineering in rice breeding will continue to grow.
Conclusion
Japan’s research in rice genetic transformation has evolved
from foundational methods to a versatile technological system, with
Agrobacterium-mediated transformation at its core. This platform
supports global eorts in trait improvement, including resistance,
quality, and nutrition. Although regulatory and public perception
barriers limit local commercialization, Japan’s innovations provide
critical support for next-generation breeding techniques. Moving
forward, combining established transformation systems with
gene editing and synthetic biology will enable more precise and
multifunctional rice improvement, contributing signicantly to global
food and nutrition security.
Funding source
National Undergraduate Innovation and Entrepreneurship
Training Program of China (202511439022)
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