Ratoon rice technology

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CHAPTER FOUR

Ratoon rice technology: A green


and resource-efficient way for rice
production
Weiqin Wangb, Aibin Hec, Guanglei Jiangc, Huijuan Sunc, Min Jiangc,
Jianguo Manc, Xiaoxia Lingc, Kehui Cuic, Jianliang Huangc,
Shaobing Pengc, Lixiao Niea,c,d,*
a
Hainan Key Laboratory for Sustainable Utilization of Tropical Bioresourse, College of Tropical Crops, Hainan
University, Haikou, China
b
Hunan Agricultural University, College of Agronomy, Changsha, China
c
MOA Key Laboratory of Crop Ecophysiology and Farming System in the Middle Reaches of the Yangtze
River, College of Plant Science and Technology, Huazhong Agricultural University, Wuhan, China
d
Hubei Collaborative Innovation Center for Grain Industry, Yangtze University, Jingzhou, China
*Corresponding author: e-mail address: 994085@hainanu.edu.cn

Contents
1. Introduction 136
2. Grain yield in the ratoon rice system 138
3. Grain quality of ratoon rice 140
4. Economics and sustainability of the ratoon rice system 142
5. The morphology and physiological bases of ratoon rice 144
5.1 Morphology of ratoon rice 144
5.2 The difference in the ratooning ability among the buds that regenerated
from different nodes of the main crop stubble 146
5.3 Nutrient translocation from the main season stubble 148
5.4 The inheritance and genetic mechanisms of the ratooning ability 148
6. Agronomic management practices to promote bud regeneration
and increase the grain yield of ratoon crop 149
6.1 Variety selection for rice ratooning 149
6.2 Establishment methods of the main season rice 151
6.3 Nitrogen application 152
6.4 Water management 154
6.5 Harvest of main crops 155
6.6 Pest and disease control 157
7. Constraints and prospects for ratoon rice production 158
7.1 Lodging 158

Advances in Agronomy, Volume 159 # 2020 Elsevier Inc. 135


ISSN 0065-2113 All rights reserved.
https://doi.org/10.1016/bs.agron.2019.07.006
136 Weiqin Wang et al.

7.2 Mechanical harvesting of main crops 158


7.3 Variety breeding for ratoon rice 159
8. Concluding remarks 160
Acknowledgments 160
References 161

Abstract
Ratoon rice, which refers to the production of a second rice crop from the stubble after
the harvest of the main crop, is considered a green and resource-efficient rice produc-
tion system. This review was undertaken to comprehensively evaluate the performance
of ratoon rice in comparison with other rice cropping systems regarding the grain yield,
grain quality, economics, and sustainability. In addition, the morphology and physiolog-
ical bases associated with the performance of ratoon rice was also reviewed. With good
crop management, ratoon rice gives a yield equivalent to 60% that of the main crop but
with a 50% reduction in resources and labor input. Moreover, rice ratooning improves
the grain quality, increases the profits for farmers, and decreases greenhouse gas emis-
sions. Several agronomic practices for tiller-bud regeneration, such as nutrient and
water management at the harvest of the main crop, establishment of the main crop,
stubble cutting height and the variety selection and integrated technologies for the
control of diseases and insects, were suggested to strongly affect the performance
of ratoon rice. However, constraints, including low and unstable yield of ratoon crop,
lodging, and lack of suitable cultivars and specialized rice harvester for the ratoon rice
system, have limited the wide scale adoption of ratoon rice. Meanwhile, the mecha-
nisms underlying the specific management strategies for ratoon rice need to be deeply
studied.

1. Introduction
Rice (Oryza sativa L.) is the second largest source of food worldwide
and has been grown on 163.2 million ha, with an average annual production
of 740.9 million tons across the globe (FAOSTAT, 2014). It has been esti-
mated that the global rice production needs to increase by 116 million tons
by 2035 to meet the rising demand for rice (Yamano et al., 2016). However,
the arable land used for rice production has been decreasing in recent years
because of urbanization and industrialization (Long, 2014), which threatens
the further increase of global rice production. The increasing multiple crop
index has been suggested to be a promising strategy to ensure total rice pro-
duction via more frequent rice harvests on existing land (Peng et al., 2009).
Generally, multiple rice cropping systems include double-season rice, triple-
season rice and ratoon rice, among which double-season rice is the dominant
Ratoon rice technology 137

multiple rice cropping system globally, especially in Asia (Yang et al., 2010).
The wide adoption of double-season rice increases the total rice production
output per land area compared to single-season rice systems and thus
contributes substantially to the global rice supply (Ray and Foley, 2013).
Nevertheless, the planting area of double-season rice has been decreasing
rapidly in recent years due to labor shortages, low degrees of mechanization,
and low production efficiency (Xu et al., 2018). To guarantee the safe pro-
duction of rice, strategies to increase the sustainability and profitability of
multiple rice cropping systems are desperately needed.
Ratoon rice, which refers to the production of a second rice crop from
the stubble left behind after the harvest of the main crop, has emerged as an
alternative option to replace double-season rice (Chen et al., 2018) (Fig. 1).
Rice ratooning is an ancient technology, which has been practiced by
farmers in China since the West Jin Dynasty (AD 265–316) (Guo, 1993).
This technology has been adopted in many countries, such as China, India,
Japan, USA, Philippines, Brazil, Colombia, Swaziland and Thailand (Faruq
et al., 2014). In the United States, ratoon rice is practiced on a commercial
scale (Harrell et al., 2009). However, in most rice production regions, rice
ratooning is only practiced by farmers as an afterthought. In recent years,
however, ratoon rice has drawn much attention and has been widely prac-
ticed by farmers because of its advantages in labor, seed, water, pesticides,
and seedbed savings. The planting area of ratoon rice was progressively
increased from 6667 ha in 2010 to 200,000 ha in 2018 in Hubei province,
China (the data were obtained from the agricultural administration of
Hubei province, China). Such a transition was mainly attributed to the sus-
tainability and high economic returns of ratoon rice in comparison with
double-season rice. It has been reported that ratoon rice gives a yield roughly
equivalent to 40–50% that of the main crop, but with a 50–70% reduction in

Fig. 1 Pictorial illustration of ratoon rice. (A) The ratoon crop at 5 days after the
harvesting of the main crop, (B) the ratoon crop at 15 days after the harvesting of
the main crop, (C) the ratoon crop at the heading stage.
138 Weiqin Wang et al.

labor and water input (Krishnamurthy, 1988; Munda et al., 2009). The costs
for seeds, fertilizers and land preparation are also largely reduced, which sub-
stantially increases the profits for farmers. Moreover, the ratoon rice system
has a shorter crop growth duration compared with double-season rice,
which is suitable to be adopted in regions where the annual cumulative
temperature is considerably higher than that required for single cropping rice
but is too low for double-season rice (Dong et al., 2017).
Research on rice ratooning started in the 1950s (Roy, 1959). Research
topics on the economics (Zhang and Gou, 1990), crop management prac-
tices (Calendacion et al., 1992; Coale and Jones, 1994; Turner and Fund,
1993) and variety selection (Kupkanchanakul et al., 1990) for ratoon rice
have been reported in the last century. However, the research progress
on rice ratooning over the last decades has not been comprehensively sum-
marized. Furthermore, the traditional ratoon rice practice with the main
crop harvested manually is no longer suitable for the current social and eco-
nomic situation due to the high labor inputs. It is important to develop a rice
ratooning technology with the main crop harvested mechanically. The pre-
sent review summarizes the latest research progress on the grain yield, grain
quality, economic input and returns, as well as the greenhouse gas emission
in ratoon rice. In addition, the morphology and physiological bases associ-
ated with the ratooning ability, the integrated agronomic technologies to
improve the productivity and sustainability of the ratoon rice system, and
the problems and prospects in ratoon rice systems are also reviewed.

2. Grain yield in the ratoon rice system


The grain yields of the ratoon crop ranged from 1.1 to 4.9 t ha 1,
which accounted for 28.6–64.3% of the grain yield in the main season
(Table 1). The large variation in the ratoon season yield among different rice
production regions might be due to the differences in the climate conditions,
varieties and crop management practices. The grain yield of ratoon crop in
tropical areas was lower than that in subtropical and temperate regions
(Olivier et al., 2014; Sinaga et al., 2014). Previously, Wang et al. (2016b)
reported that the high temperature in tropical areas reduced the radiation
use efficiency and resulted in a yield reduction compared with temperate
and subtropical areas. The high annual temperature might have shortened
the rice growth duration and thus limited the yield potential of ratoon rice.
However, the sufficient accumulated temperature in tropical areas allows the
practice of tropical perennial rice (SALIBU), which refers to the harvest of
Ratoon rice technology 139

Table 1 Main season yield, ratoon season yield and the ratio between ratoon and main
season yields in different rice producing regions.
Grain yield
(t ha21)
Main Ratoon Ratio between ratoon season yield
Location season season and main season yield (%) References
The 9.7 3.3 34.0 Dou et al.
United (2016)
States
Nigeria 6.5 2.9 44.6 Adigbo et al.
(2012)
Indonesia 4.4 2.0 45.5 Susilawati
et al. (2010)
Korea 6.1 2.3 37.7 Shin et al.
(2015)
Indonesia 2.6 1.2 46.1 Sinaga et al.
(2014)
Vietnam 4.2 2.7 64.3 Sen and
Bond (2017)
West 4.2 1.2 28.6 Olivier et al.
Africa (2014)
China 9.0 4.9 54.4 Dong et al.
(2017)

ratoon crops 3.5–4 times annually and realizes an equivalent or a higher yield
in ratoon crop than the main crop under the tropical, winterless climate
(Pasaribu and Anas, 2018). It has been reported that with the multiple har-
vest of the ratoon crops for SALIBU, the ratoon yield can achieve 5–9 t ha 1,
which was even higher than that of the main season crop (Yamaoka et al.,
2017). In subtropical and temperate regions, a ratoon season yield of
3–6 t ha 1 can be achieved with the appropriate variety selection and good
crop management (Dong et al., 2017; Dou et al., 2016; Oad et al., 2002).
A relatively higher ratoon crop yield of 9.7 t ha 1 was observed when suit-
able rice varieties were incorporated and the best management practices
were applied (Li et al., 2009). Nevertheless, in temperate regions, if the sow-
ing date of the main season rice was postponed or the late ripening varieties
were adopted for rice ratooning, the ratoon crop might not mature due to
the low temperature during the grain filling stage. Therefore, it is necessary
140 Weiqin Wang et al.

to develop rice varieties that are cold tolerant and those with a relatively
shorter growth duration (7–15 days shorter than the currently adopted
ratoon rice varieties) (Dong et al., 2017). Other factors that influence the
grain yield of ratoon rice include the planting date and establishment method
of the main crops, stubble cutting height, fertilizer and water management,
which will be fully discussed in the following sections.

3. Grain quality of ratoon rice


The quality of rice, which is evaluated based on several aspects includ-
ing milling, appearance, cooking, and nutrient qualities, has drawn more and
more attention from consumers because of the growing economy and
increased purchasing power of a considerable part of the population
(Fitzgerald et al., 2009). Ratoon rice is characterized as having high grain
quality, which could be a determinant factor in the economic returns for
farmers (Han et al., 2004; Yazdpour et al., 2012). The improved grain qual-
ity of ratoon rice has been widely reported previously (Fig. 2). Research
by Alizadeh and Habibi (2016) reported that rice ratooning improved the
milling and appearance quality of rice. Compared with that of the main crop,
the head rice yield of the ratoon crop was increased by 5.5%, while the
chalkiness degree was decreased by 10.9% (Fig. 2A and B). The improved
nutrient, cooking and taste quality of the ratoon crop, which was reflected
by the increased amylose content (Fig. 2C), decreased gelatinization temper-
ature and gel consistency of the rice grains (Fig. 2D and F), was also
observed, compared with that of the main crop. In addition, the grain quality
of the ratoon crop was superior to that of the double-season rice (Fig. 2G–J).
When compared with the early crop, the head rice yield, milled rice yield
and total starch content of ratoon crop were significantly increased by 2.0%,
6.3% and 3.9%, respectively. Moreover, those attributes were increased by
1.4%, 0.4% and 2.7%, respectively, compared with that of the late-season
rice (Fig. 2F and G). Moreover, rice ratooning decreased the chalkiness
by 10.1% and 6.2%, compared with that of early and late-season rice, respec-
tively (Fig. 2I). The better grain quality of ratoon rice might be due to the
lower accumulative temperature during the growth stage of ratoon rice, in
comparison with the main crop or double-season crops. A lower tempera-
ture extended the grain maturity period, promoted the process of grain
filling, and thus resulted in better milling, appearance, and cooking and taste
quality in ratoon rice (Funaba et al., 2006). Our latest research revealed that
rice ratooning increased the phenolic compounds, lipids and lysine content
Ratoon rice technology 141

Fig. 2 The comparisons of grain quality between main season rice and ratoon season
rice, and between ratoon season rice and double-season rice. (A–E) The comparisons of
head rice yield, chalkiness rate, amylose content, gel consistency and gelatinization
temperature between main season rice and ratoon season rice, (F–I) the comparisons
of head rice yield, chalkiness rate and total starch content among ratoon season rice,
early-season rice and late-season rice.

in the rice grains (unpublished data). The mechanisms associated with the
improved nutrients content in ratoon crop remain unknown. However,
one of the hypotheses is that ratoon rice might have higher carbohydrate
and nitrogen transportation efficiency than single rice crops, which pro-
motes the synthesis of phenolic compounds, lipids and lysine. Nevertheless,
several studies reported that rice ratooning did not increase or decrease the
grain quality of rice compared with that of the main crops in tropical regions
(Bollich and Turner, 1988; Shin et al., 2015). The high temperature during
142 Weiqin Wang et al.

the grain filling stage might be one of the reasons that contributed to the
quality decrease in ratoon rice. A high temperature during kernel develop-
ment caused spikelet infertility, reduced the grain weight and yield, resulted
in a greater proportion of chalky grains and affected the grain physicochem-
ical attributes by lowering the amylose content (Cooper et al., 2008; Kadam
et al., 2014; Peng et al., 2004). In addition, lodging during the main season,
as well as the rolling damage caused by the harvest machine, might result in
the poor grain quality in the ratoon season.

4. Economics and sustainability of the ratoon rice


system
Rice ratooning reduces the process of land preparation, sowing and
transplanting, which minimizes the cost for labor, water and seeds. It has
been reported that the labor and seeds input for the ratoon rice system
was reduced by 28.5% and 51.9%, respectively, compared with double-
season rice system (Liang et al., 2016; Sen and Bond, 2017). The cost for
fertilizers and pesticides in ratoon systems was also reduced by 31.5% and
41.1%, respectively, which ultimately decreased the total input by 31.1%
compared with that of double-season cropping (Fig. 3). Although the annual
yield of the ratoon rice system was lower than that of double-season rice, the
net profit was higher than that of double-season rice (Table 2). Munda et al.
(2009) reported that the profits in the ratoon rice system increased by
19 USD/ha, which was mainly attributed to the significant decreases in
the cost for labor, fertilizers and seeds, compared with double-season rice
(Table 2). Meanwhile, compared with single-season rice, the profits in
the ratoon rice system increased by 125.4% (Table 2). When evaluating
the economic benefits between ratoon season rice and the early and late
crops in double-season rice system, the profit in ratoon rice was lower than
that of the late crop but was much higher than that of the early crop
(Table 2). This was mainly because the grain quality of ratoon rice was better
than that of the early crop, which minimized the return gap and increased
profits. In addition, Firouzi et al. (2018a) demonstrated that the total energy
consumption for producing 2.4 kg of brown rice as the feedstock unit for 1 L
bioethanol production was lower in the ratoon crop than in the single crop,
thereby indicating that the bioethanol feedstock production in ratoon
rice was more energy efficient than in the single rice cropping system.
In addition, in South and Southeast Asia, flooding stress greatly impacts
the rice production and could cause 43.75% of yield loss per year
Ratoon rice technology 143

Fig. 3 The comparisons of the seed, fertilizer, labor, pesticide and total costs between
the ratoon rice system and double-season rice system. Data resources: Sen, L.T.H. and
Bond, J., 2017, Agricultural adaptation to flood in lowland rice production areas of central
Vietnam: understanding the ‘regenerated rice’ ratoon system, Clim. Dev. 9, 274–285;
Liang, Y., Zhang, Q., Zhou, J., Li, Y., Tan, C., Huang, H., 2016. Effects of different cultivation
patterns on rice quality and economic efficiency. Acta Agr Boreali Sinica. 31, 265–269.
(In Chinese with English Abstract).

Table 2 Costs, returns and profits of the ratoon rice system in comparison with
single-season rice and double-season rice.
Cost Returns Profit
Location Cropping system USD/ha USD/ha USD/ha References
India Double-season rice 343 616 274 Munda et al. (2009)
Rice-ratoon rice 271 564 293
Single-season rice 205 335 130
China Early-season rice 1703 2415 712 Liang et al. (2016)
Late-season rice 1740 2620 880
Ratoon rice 509 1439 864

(Sen and Bond, 2017). Meanwhile, rice ratooning was practiced as an auton-
omous adaptation by farmers to overcome the economic losses from the
main season crops caused by flooding, as the profit of rice ratooning was sim-
ilar or even higher than that of single rice cropping (Manzanilla et al., 2011).
144 Weiqin Wang et al.

Rice paddies are the major source of CH4 and N2O and can also be a
source or sink of CO2, which contributed to approximately 30% and
11% of global agricultural CH4 and N2O emissions, respectively (Hussain
et al., 2015; Jat et al., 2016; Tao et al., 2016). Therefore, it is necessary to
evaluate the sustainability of ratoon rice in comparison with other rice
cropping systems. Firouzi et al. (2018b) evaluated the performances of the
ratoon rice and single rice systems regarding greenhouse gas emission.
The results showed that the CO2, N2O, and CH4 emissions from the ratoon
rice system were less than that from the single rice cropping system. Similar
results were also observed by Lindau and Bollich (1993). The beneficial
effect of rice ratooning in reducing greenhouse gas emission was mainly
attributed to the lower fossil fuel consumption, middle season drainage
and decreased organic matter content compared with single rice crops
(Firouzi et al., 2018a). In summary, the above studies indicated that ratoon
rice is an economic, green and resource-efficient cropping system, which
could increase the sustainability of rice production system. However, a series
of studies is required to evaluate the sustainability of ratoon rice on a
comprehensive scale.

5. The morphology and physiological bases of


ratoon rice
The ratooning ability refers to the potential of the dormant buds to
regenerate from the nodes of the stubble, which is fundamental to the yield
formation of ratoon season rice. The regeneration ability is a complex trait
that is dependent on many heritable and environmental factors. Therefore,
understanding the morphology and physiology traits of ratoon rice in asso-
ciation with the ratooning ability is essential for further improvement of
ratoon rice. The present section introduces the morphology traits of ratoon
rice regarding the growth of its auxiliary buds, leaf, stem and root. In addi-
tion, the differences in the ratooning ability among different nodes, the
influence of nutrient translocation on the ratooning ability, and the inher-
itance and genetic mechanism associated with the ratooning ability are also
fully discussed.

5.1 Morphology of ratoon rice


The morphology traits of ratoon season rice are quite different from that of
the main crop (Fig. 4). The life cycle of ratoon season rice starts with the
sprout of dormant buds from the nodes of the stubble. The dormant buds
Ratoon rice technology 145

Fig. 4 The diagrams of the nodes and regenerated panicles in the ratoon rice system.
The ratoon tillers were regenerated from the nodes of the main crop stubble.

start to sprout during the reproductive stage of the main crops (Wang and
Tang, 1991). However, at this stage, the growth speed of the ratoon buds
is very slow because most of the nutrients are transported to the panicles of
the main crops, which limits the growth of the auxiliary buds (Huang and
Zhang, 1989). After the dough ripening stage of the main season crops, the
sprout speed of the auxiliary buds is significantly increased, and the ratoon til-
ler numbers reach the maximum level at 15–20 days after the harvesting of the
main crops (Huang and Zhang, 1989). Meanwhile, panicle differentiation of
the ratoon buds starts at 15 days after the heading stage of the main crops (Lin
et al., 2015). In general, it takes 40–60 days for the ratoon buds to finish panicle
differentiation (Huang and Zhang, 1989). Significant variations in the growth
of ratoon buds from different nodes of the stubble have been compared, and
the auxiliary buds from the basal nodes sprouted earlier than that from the
upper nodes, but the upper auxiliary buds are suggested to have a faster growth
speed than the basal ones (Prashar, 1970).
The process of heading and grain filling for ratoon crop mainly relies on
the photosynthesis products, which are transported from the leaves,
suggesting the important roles of ratoon leaf on the growth and yield in
ratoon season. The leaf number of the ratoon crop is less than that of the
main crop. It has been reported that the leaf area index (LAI) for the ratoon
crop was generally approximately 1–2, which was much lower than that of
146 Weiqin Wang et al.

the main crops (Zhang et al., 2002). The lower leaf number and LAI in the
ratoon crop were mainly attributed to the shorter vegetative growth stage of
the ratoon crop compared with the main crop. The lower root growth dur-
ing the ratoon season was also suggested to be responsible for less leaf areas
and leaf numbers for the ratoon crop (Zhang, 1995). However, the smaller
leaf area resulted in a smaller shading area and K value for the ratoon crops
(Zhang, 1995). It has been reported that the photosynthetic rate in the
ratoon crop was higher than that of the main crop, which is beneficial
for the accumulation and transportation of photosynthesis products
(Zhang, 1995).
The stem of the ratoon crop is comprised of the main stems from the
stubble of the main crop, and new stems developed from the auxiliary buds.
Comparing with the main crop stem, the ratoon crop stem is shorter and
thinner with less nodes number and less internodes elongation (Chen
et al., 2010). Moreover, the stems that regenerated from the basal nodes
of the main crop stubble are longer and thicker than those regenerated from
the upper nodes (Chen et al., 2010).
The roots of ratoon rice include newly developed root and the old roots
from the main crop. It has been suggested that the old roots from the main
crops played major roles in supporting the growth of ratoon crops (Chauhan
et al., 1985), which implied that the limited growth of ratoon roots might be
a constraint to the further increase in the regeneration ability. Generally, the
newly developed roots in the ratoon season are regenerated from the nodes
of the main crop stubble or from the nodes of the newly developed ratoon
tillers. The growth of new roots in ratoon crops can be regulated by the stub-
ble height. Low cutting height of the stubble could significantly promote the
growth of new roots in ratoon season. The root vigor of ratoon crops was
also suggested to be influenced by the fertilizer management and the above-
ground biomass of the main crop (Hillers, 1988; Muhammad et al., 2008).
However, the relationship between new roots and old roots in regulating the
regeneration ability of ratoon crops still needs to be further developed.

5.2 The difference in the ratooning ability among the buds that
regenerated from different nodes of the main crop stubble
In ratoon season, there are generally three to seven nodes at the stubble that
can regenerate panicles and contribute to the grain yield of the ratoon crop.
The ratooning ability of the auxiliary buds varied among different nodes
of the stubble (Chauhan et al., 1985). It has been reported that the sprout
of the auxiliary buds started from the upper nodes of the stubble, and the
regeneration rate was decreased from upper nodes to the basal nodes
Ratoon rice technology 147

(Sun et al., 1988). The yield components of ratoon season rice also varied
between the panicles that regenerated from the upper nodes or the basal
nodes. The upper nodes produced more panicles than the basal nodes, while
the panicles from the basal nodes were large than the upper ones, but the
fertility percentage of the spikelet regenerated from the basal nodes was
lower than that of the upper nodes (Sun et al., 1988). The variance in the
yield composition between the upper nodes and the basal nodes on resulted
in different contributions to the grain yield of the ratoon crop. According to
Dong et al. (2017), the panicles regenerated from the second nodes from the
top contributed to 66.6% of the ratoon season yield, which was mainly
attributed to the increased panicle numbers and the higher filled grain rate,
compared with that of basal panicles (Table 3). In contrast, Yuan and Sun
(1996) observed that the yield formation of ratoon crop was mainly attrib-
uted to the panicles that regenerated from the basal nodes rather than the
upper nodes (Table 3). The mixed results among different studies might
be caused by the difference in the varietal traits. The rice varieties used
for ratooning can be classified into “high stubble type” and “low stubble
type.” For “high stubble type” varieties, the second and third nodes from
the top have the strongest regeneration ability and contributed to almost
90% of the ratoon season grain yield (Yi et al., 2008), while for “low stubble
type,” the basal nodes are the dominant nodes to regenerate panicles, which
contributed to most of the grain yield in ratoon season (Cai et al., 1998;
Chen et al., 1996). Further research is needed to fully understand the factors
that affect the ratooning ability among the buds, which are regenerated from
different nodes of the main crop stubble.

Table 3 The yield components and contributions to the total grain yield among
panicles regenerated from different nodes in the ratoon season.
Filled 1000-grain Yield
Panicles grain rate weight contribution
Spikelets
Position No m22 panicles21 (%) (g) (%) References
D2 283 a 69.0 a 76.7 a 20.0 a 66.6 Dong et al.
(2017)
D3 196 b 54.0 ab 62.5 b 20.6 a 30.5
D4 46 c 39.2 b 40.1 c 18.6 a 2.9
D2 71 36.9 79.3 24.1 33.8 Yuan and
Sun (1996)
D3 96 39.8 72.9 22.9 42.3
D4 66 39.8 60.1 22.6 23.9
148 Weiqin Wang et al.

5.3 Nutrient translocation from the main season stubble


At the early growth stage of the ratoon crop, the sprout of ratoon tillers and
the differentiation of panicles were mainly attributed to the nutrient stored
in the rice straws of the main crop (Bollich et al., 1980). Moreover, the
nutrient stored in the rice straws was positively related with the regeneration
ability of the ratoon crop (Chen et al., 2018; Santos et al., 2003). Non-
structural carbohydrate and nitrogen are the major components of the nutri-
ent stored in the rice straw (Balasubramanian et al., 1970). The ratio between
carbohydrate (C) and nitrogen (N) in the straws influences the regeneration
ability of ratoon crops (Vergara et al., 1988). It was reported that the
N content in the main crop straw was negatively correlated with the
ratooning ability in ratoon season (Vergara et al., 1988), while the carbohy-
drate in the rice straw was positively related to the ratooning ability (Dat and
Peterson, 1983). In addition, it was documented that the source and sink
relationship in the main season regulated the nutrient accumulation in rice
straws and then influenced the regeneration of the ratoon crop (Xu et al.,
2015). Vigorous source activity promoted the transportation of photosyn-
thesis products from the leaves to the rice straws, and thus accelerated the
sprout of ratoon buds (Ren et al., 2006). By contrast, early leaf senescence
decreased the nutrient accumulation in rice straws and decreased the
ratooning ability in ratoon season (Cuevas-Perez, 1980). Meanwhile, the
sink size of the main crop negatively affected the regeneration rate of ratoon
crop (Xu et al., 1997b). For varieties with a larger sink size, more photosyn-
thates would be transported to the panicles of the main crop, which in turn
decreased the nutrient accumulation in rice straws, compared with varieties
with a smaller sink size (Xu et al., 1997a). The leaf/grain ratio of the main
crop was also negatively related with the nutrient accumulation in the rice
straw, suggesting that the leaf/grain ratio might be used as an indicator to
predict the regeneration ability of ratoon crop (Xu et al., 2000). However,
the quantitative relationship between the source-sink in the main crop and
the ratooning ability of the ratoon crop have not been established and still
need to be clarified in future studies.

5.4 The inheritance and genetic mechanisms of the ratooning


ability
The ratooning ability is a complex trait, which is determined by heritable
and environmental factors (Hillers, 1988). Understanding the inheritance
and genetic mechanisms associated with the ratooning ability is necessary
Ratoon rice technology 149

for the breeding of high ratooning ability varieties. Jiang (1996) suggested
that the ratooning ability was controlled by a single gene. However, most
of the previous studies reported that the ratooning ability was controlled
by multiple genes (Li and Chen, 1988; Lin et al., 2015). Li and Chen
(1988) observed significant cytoplasmic effects and nucleo-cytoplasmic
interaction for the inheritance of the ratooning ability. Chauhan et al.
(1989) suggested that additive effects were more important than the dom-
inance effects for the ratooning ability. Moreover, this research implied that
the interaction effect between genomic and environment was appreciable in
early generations but was less pronounced in advanced generations. In recent
years, QTL mapping for the major traits concerning the ratooning ability of
rice has been conducted by several researchers (Mao and Cheng, 1999; Yang
et al., 2012). Yang et al. (2012) detected two QTLs that affected the
ratooning ability on chromosomes 4 and 5; further analysis revealed that
the contribution rates of these two QTLs were 8.17% and 7.19%, respec-
tively. Zheng et al. (2004) reported that three QTLs conferring the
ratooning ability (ratoon panicle number), seed setting rate and yield of
the ratoon crop were all located on RMl8-RM234 of chromosome 7 and
had a lower contribution rate. Meanwhile, five QTLs controlling the head-
ing date, plant height, spikelets per panicle, panicle length and density of the
main crop were all mapped on RML-RM283 of chromosome 1, and had a
higher contribution rate for panicle traits. However, the inheritance and
genetic mechanisms associated with the ratooning ability have not been fully
explored.

6. Agronomic management practices to promote bud


regeneration and increase the grain yield of
ratoon crop
The optimum agronomic practices in both the main and ratoon sea-
sons are crucial to guarantee the sustainability and productivity of ratoon rice
system. The present section summarizes the high yielding technologies for
the ratoon rice system, including the variety selection, establishment and
harvest methods, water and nitrogen management and pest and disease
prevention (Fig. 5).

6.1 Variety selection for rice ratooning


Up to now, there has been no specific breeding project for ratooning rice.
Therefore, the varieties used in ratoon rice systems are screened out from the
150 Weiqin Wang et al.

Fig. 5 Demonstration of integrated agronomic technologies for the ratoon rice system.
The technologies include variety selection, nitrogen management, water management,
incorporation of mechanical harvesting and direct-seeding, stubble cutting height and
disease, pest and lodging control.

varieties that have been widely popularized in production. Several varietal


traits, such as growth duration, regeneration ability and the yield potential,
should be considered when evaluating the suitability of varieties to practice
as ratoon rice. The growth duration is generally positively correlated with
the yield potential of a variety. The intermediate- to late-maturing varieties
produce higher ratoon yields than the early-maturing varieties (Bahar and
De Datta, 1977; Krishanamurthy, 1988). However, in temperate and sub-
tropical regions, the selection of long growth duration varieties may lead
to the risk of yield loss due to the limited annual accumulated temperature.
However, the varieties with short growth durations were suggested to have a
low yield potential, which could decrease the profits and production effi-
ciency for farmers (Khush, 1995).
The ratooning ability is a potentially important and complex trait, which
is variable among different types of varieties. Previously, Chen et al. (2018)
reported that the regeneration in hybrid varieties was higher than in inbred
varieties, and the higher regeneration rate was associated with the higher dry
weight per stem after harvesting of the main crop in hybrid rice cultivars
compared with the inbred ones. Susilawati et al. (2010) suggested that the
new plant type variety had better vegetative growth and higher ratoon sea-
son yield than other types of varieties. Lin et al. (2015) recommended that
two lines hybrid varieties generally had higher ratooning performance than
three lines hybrid varieties. In Malagasy, japonica varieties have a better
ratooning ability than indicas, and japonica/indica crosses have shown high
Ratoon rice technology 151

ratooning ability (Krishanamurthy, 1988). Several indices of the main season


have been suggested for use as indicators for the high regeneration ability of
ratoon crops. It has been indicated that the stem thickness of the main crop
can reflect the regeneration ability of ratoon crop (Hillers, 1988; Ichii and
Sumi, 1983). The ratio between leaf and grains during the heading stage
of the main crops has also been suggested to be a good indicator for the
ratooning ability (Xu et al., 2000). However, so far, no definite indicators
closely related to a strong ratooning ability have been found.

6.2 Establishment methods of the main season rice


Direct-seeded rice is a simplified and sustainable rice establishment method,
which can potentially reduce the consumption of resources and decrease the
emissions of greenhouse gases while maintaining grain yields compared with
transplanting (Kumar and Ladha, 2011; Tao et al., 2016). Incorporating
direct-seeded rice into the ratoon rice system to develop direct-seeded
ratoon rice (DSR-RR) is a promising strategy for the sustainable develop-
ment of the ratoon rice system. Previously, Dong et al. (2017) found that
there were no significant differences in the ratoon season yield and annual
yield between direct-seeded ratoon rice and transplanted ratoon rice systems
(Table 6). Nevertheless, there is a potential for DSR-RR to replace
TR-RR as an alternative option. Nevertheless, the reduced grain yield in
DSR-RR in both the main and ratoon seasons was also observed by several
researchers (Liu et al., 2012; Yang et al., 2017) (Table 4). Several factors,
including poor and uneven crop establishment, inadequate weed control
and lodging susceptibility, in direct-seeded rice might be responsible for
the decreased grain yield in DSR-RR (San-Oh et al., 2004). In addition,
the growth duration of DSR-RR in the main field is longer than that of

Table 4 The comparisons of main season yield, ratoon season yield and annual growth
duration between direct-seeded and transplanted ratoon rice.
Main season Ratoon season Annual growth
yield yield duration
Establishment
21 21
method t ha t ha (days) References
Direct-seeding 9.2 a 4.8 a 196 Dong et al.
(2017)
Transplanting 8.7 a 5.1 a 198
Direct-seeding 6.8 b 3.0 b 185 Yang et al.
(2017)
Transplanting 8.1 a 3.5 a 196
152 Weiqin Wang et al.

TR-RR due to the lack of a nursery stage. If the sowing date of the main
crop was postponed, the ratoon crop would not mature because of low tem-
peratures during the grain filling stage. Strategies to improve the yield sustain-
ability of DSR-RR have been proposed, including developing effective pre-
sowing seed treatment, such as seed priming and seed coating, to enhance the
seedling establishment of direct-seeded rice under adverse environment (Mei
et al., 2017; Wang et al., 2016a; Zheng et al., 2016), prevent the infestation of
weeds and weedy rice (Chauhan, 2013), breed and select varieties with high
lodging and cold tolerance (Hyun et al., 2017; Liu et al., 2015) and develop
sowing machines for direct-seeded rice (Wang et al., 2017).
The planting date of the main crops should be dependent on the rainfall
and temperature during the growth periods. For instance, in East Asia, early
planting may lead to chilling stress to the main season crops, which could
result in poor seedling establishment (Sipaseuth et al., 2007). If the planting
date of the main crops is delayed, the ratoon crops would not be able to
mature due to the low temperature during the grain filling stage
(Shimono et al., 2007). In South and Southeast Asia, the planting date of
the main crops should be based on the frequency of the rainfall. Heavy rain-
fall during the reproductive stage of rice would result in severe yield loss
(Wassmann et al., 2009).

6.3 Nitrogen application


Nitrogen (N) was reported to have the most significant effects on the growth
and grain yield of ratoon crops (Mengel and Leonards, 1978). The positive
effects of N on improving tillering, increasing the sprouting rate of the buds
from the lower nodes of the stubble, promoting the grain filling and increasing
the grain yield of the ratoon crop have been reported (Sun et al., 1988; Xu
et al., 2006). In the main season, the N rate for the rice crop should be less
than that of the local high yielding practice for single rice system to reduce
the risk of lodging and disease infection, and it was recommended that splitting
3 times with 35% of N should be applied as a starter basal dose, while the resid-
ual N should be equally split at the middle tillering stage and the panicle ini-
tiation stage of the main crop (Islam et al., 2008; Liu et al., 2014).
To promote the regeneration of ratoon buds and increase the produc-
tivity of ratoon rice, two additional N fertilizations are needed. First, the
bud-promoting fertilizers were strongly recommended to be applied at
approximately 15 days after heading in the main season to improve the
regeneration of ratoon buds (Nakano and Morita, 2008). The application
Ratoon rice technology 153

of N during this stage increased the photosynthetic ability of flag leaves, pro-
moted the nutrient transportation to the stems and activated the growth of the
ratoon root system, which largely promoted the sprout of regenerated buds
and thus increased the grain yield of ratoon crops (Nakano and Morita,
2008; Xu et al., 1997a,b). Meanwhile, the early application of bud-promoting
fertilizers increased the risks of lodging for the main crop (Chauhan et al.,
1989; Xu et al., 2000). The optimum N rate for the bud-promoting fertilizer
varied across different studies (Table 5). It has been observed that the grain
yield of ratoon crop was positively correlated to the rate of bud-promoting

Table 5 The effect of nitrogen application rate on the grain yield of ratoon crop.
N rate Ratoon crop yield
Location kg ha21 t ha21 References
The United States 30 2.4 c Bond and Bollich (2006)
60 2.6 b
90 2.7 a
120 2.8 a
Iran 0 1.1 c Petroudi et al. (2011)
35 2.5 b
70 2.8 a
Iran 0 0.6 c Huossainzade et al. (2011)
30 0.6 c
60 1.4 a
90 1.0 b
China 0 5.3 Yu et al. (2017)
70 5.5
95 5.7
115 5.7
140 5.3
160 5.0
180 4.8
205 4.6
154 Weiqin Wang et al.

fertilizer in the range of 0–120 kg N ha 1 (Bond and Bollich, 2006; Petroudi


et al., 2011) (Table 5). However, excess N input for the bud-promoting fer-
tilizers would further decrease the grain yield of the ratoon crops
(Huossainzade et al., 2011). The above research suggested that quantification
of the rate of bud-promoting fertilizers is necessary. It has been reported that
the N requirement for ratoon crop was negatively correlated to the leaf
N content indirectly reflected by the leaf SPAD value at the heading stage
of the main crop (Xu et al., 1997a; Yu et al., 2013). However, the quantitative
relationship between the rate of bud-promoting fertilizers and the leaf SPAD
value of the main crop has not been determined.
Second, the application of tiller-promoting fertilizers at 2–5 days after the
harvesting of the main crop significantly improved the tillering ability and
the grain yield of ratoon crop (Turner and McIlrath, 1988). Increasing
the rate of tiller-promoting fertilizers in the range of 0–90 kg N ha 1 signif-
icantly increased the grain yield of ratoon crop (Mengel and Wilson, 1981).
However, it has also been reported that the increase of N rate from
50 to 90 kg N ha 1 did not influence the grain yield of the ratoon crop
(Nassiri et al., 2011). According to our unpublished data in subtropical
regions, the recommended N rate for tiller-promoting fertilizers is
52.5–70 kg N ha 1 to achieve the highest grain yield in the ratoon season.
Meanwhile, the application of potassium fertilizer (K) for the ratoon crop
was based on the soil indigenous K supply and K rate applied in the
main season.
The above reports suggested that 3–5 times of fertilization are essential in
the whole growth duration of ratoon rice. Nevertheless, the time of fertil-
ization must be reduced due to labor scarcity and high labor cost, as well as
the difficulty of fertilization at approximately 15 days after heading in the
main season. In China, a specialized fertilizer for ratoon rice named “Dao
Sun Wang” has been developed. This specialized fertilizer is slow releasing,
root-protecting, and bud-promoting, which reduces the fertilization times,
improves fertilizer use efficiency, and increases the grain yield of
ratoon crop.

6.4 Water management


Soil drying is the most important water management practice to increase and
sustain the grain yield of ratoon crops. Soil drying during the maximum til-
lering stage of the main crop increased the oxygen content and oxidation-
reduction potential of the soil, which is beneficial for the growth of the root
system (Hasegawa and Yoshida, 1982). It has been reported that heavy
Ratoon rice technology 155

drying at the tillering stage increased the root activity, root wound flow, root
number and root growth ability by 20.6%, 9.94%, 11.11% and 19.25%,
respectively, compared with flooding irrigation (Lin et al., 2015). The vig-
orous root growth contributed to the regeneration of ratoon buds for ratoon
crops. In addition, soil drying during the middle-late grain filling stage of the
main crop increased the soil hardness degree, which could mitigate the
rolling damage to the ratoon crops caused by the harvest machine. Com-
pared the slight soil drying, heavy soil drying decreased the rolling damage
to the ratoon crops by 8% (unpublished data). However, too heavy drying
during the maturity stage of the main crop would reduce the ratoon season
yield by depressing the panicle differentiation of the ratoon crops. The grain
yield of the ratoon crop was reported to significantly decrease when the soil
water content during the maturity stage of the main crop was below 21%
( Jiang et al., 2005).
In the ratoon season, flooding irrigation immediately after the harvesting
of the main crops effectively promoted the tillering of the ratoon crops
(Mengel and Wilson, 1981). However, the time and amount of irrigation
should be strictly controlled because heavy and continuous flooding would
suppress the growth of the ratoon crop by decreasing the growth of roots and
the buds from the basal nodes, which could result in a 69% yield decrease in
the ratoon season compared with that of shallow flooding (Bahar and De
Datta, 1977). In summary, soil drying during the maximum tillering stage
and the middle-late grain filling stage of main crops, along with flooding irri-
gation immediately after the harvest of the main crops, are recommended to
be the efficient water management strategies for ratoon rice. Further studies
should be focused on the technologies to improve the water use efficiency in
the ratoon rice system.

6.5 Harvest of main crops


The main season rice can be harvested manually or mechanically. The man-
ual harvesting of the main crop decreased the production efficiency of the
ratoon rice system (Cai and Chen, 2000; Liu et al., 2014). Therefore,
mechanical harvesting of the main crop is the only method for the sustain-
able development of rice ratooning. However, several problems, including
decreased grain yield and grain quality in the ratoon season caused by the
rolling damage from the harvest machine, and inappropriate crop manage-
ment for mechanically harvested ratoon rice, limited the further extension of
mechanically harvested ratoon rice. These issues and the probable solutions
will be fully discussed in Section 7.
156 Weiqin Wang et al.

Table 6 The effect of stubble cutting height on the grain yield of ratoon crop.
Stubble
height Ratoon season yield
Location (cm) t ha21 References
The United States 20 2.5 Harrel et al. (2009)
40 2.3
Korea 10 2.8 a Shin et al. (2015)
20 2.1 b
30 1.9 b
40 2.2 b
Iran 10 0.5 Huossainzade et al. (2011)
25 1.0
30 1.1
China 10 2.8 Jiang et al. (2005)
25 4.0
40 5.5

The optimum stubble cutting height of the main crop was varied
depending on the ratooning characteristics of the varieties and the meteo-
rological conditions of the rice production regions (Table 6). For varieties
with a strong regeneration ability from the basal nodes of the stubble, the
main crop should be harvested at a low cutting height (Harrell et al.,
2009; Shin et al., 2015) (Table 6). By contrast, increasing the stubble cutting
height significantly increased the grain yield of the ratoon crop for the vari-
eties that mainly depended on the panicles regenerated from the upper nodes
(Huossainzade et al., 2011). Research by Jiang et al. (2005) reported that a
high stubble height (40 cm) increased the panicle numbers and filled grain
rate of the ratoon crops, and thus increased the grain yield of the ratoon crop
by 96.4% compared with a low stubble height (10 cm) (Table 6). The
meteorological conditions of the rice production regions strongly affected
the regeneration ability of the ratoon crop in response to different stubble
cutting heights. In tropical areas, low stubble cutting height in ratoon rice
(including SALIBU rice) was usually adopted by farmers (Pasaribu and
Anas, 2018). For a lower stubble cutting height, the sufficient temperature
and solar radiation not only ensure the regeneration of basal auxiliary buds
Ratoon rice technology 157

but also promote the growth of tillers regenerated from the root nodes. In
addition, a low stubble cutting height removed the apical dominance from
the upper nodes to the basal nodes and root nodes, which greatly increased
the regeneration rate and grain yield of the ratoon crop. For the ratoon crop
grown in subtropical and temperate regions, a high stubble cutting height
was recommended because the sprout of the auxiliary buds started from
the upper nodes of the stubble, and the regeneration rate decreased from
the upper nodes to the basal nodes (Sun et al., 1988). In addition, a lower
stubble cutting height would extent the growth duration of ratoon crops
in subtropical and temperate regions (Harrell et al., 2009).

6.6 Pest and disease control


The infection of diseases and pests significantly influence the productivity of
the ratoon rice system (Santos et al., 2003). It has been reported that if the main
crop suffered from systemic disease infection, the ratoon crop would be
infected by the same disease, and the stubble was unlikely to produce any
ratoon tillers (Mew and Fabellar, 1988). Meanwhile, the occurrence of pests
and diseases in the ratoon season was quite different from that in the main sea-
son. Compared with the main crop, the ratoon crop was more attractive to
leaf folder but was a poor host for caseworm. Moreover, diseases such as yel-
low dwarf, grassy stunt and stem rot more frequently occurred in the ratoon
season rather than in the main season (Muniyappa et al., 1988; Nagaraju et al.,
1988). Therefore, it is suggested that the strategies to control diseases and pests
for the ratoon crop should be different than that for the main crop.
The use of insecticides and pesticides successfully prevents and controls the
infection of pests and diseases; however, intensive pesticide use may cause
environmental contamination and increase the risk of insecticide and pesticide
resistance (Heap, 2012). As alternatives, several nonchemical methods are rec-
ommended to help reduce the pest and disease pressure without environmen-
tal pollution. The selection of varieties with high pest and disease resistance is
necessary for the successful production of ratoon rice. In ratoon season, shal-
low flooding and alternative wetting and drying water management reduced
the occurrence of stem rots in the ratoon crop (Santos et al., 2003). In addi-
tion, priming of the main crops by using methyl jasmonate (MeJA) was
suggested to enhance the anti-herbivore resistance of the ratoon crop
(Ye et al., 2017). Adjusting the nitrogen application rate and soil drying during
the middle tillering stage could effectively prevent the main and ratoon crops
from the infection of pest and diseases (Liu et al., 2014).
158 Weiqin Wang et al.

7. Constraints and prospects for ratoon rice production


7.1 Lodging
Lodging is one of the most serious problems that influence the stability of the
ratoon rice system (Kano and Kubota, 1990; Kashiwagi et al., 2005). Severe
lodging not only decreases the grain yield and grain quality of main crops but
would also drastically decrease the regeneration ability and grain yield of
ratoon crops. Chen et al. (2010) reported that lodging during the main sea-
son decreased the ratoon season yield by 30.7%. The lodging of the main
crop caused invertible damage to the stubble and roots, which limited the
nutrient supply to the ratoon buds and ultimately led to the decreased ratoon
yield. The selection of high lodging resistance varieties is one of the effective
approaches to reduce the risk of lodging in the ratoon rice system. The char-
acteristics of lodging resistant varieties are related to the crop height, the
height of the center of gravity, internode flexure resistance and thickness
of the stem and sheath (Chen et al., 2018). The ratooning ability was also
suggested to be positively correlated with the lodging resistance of rice crops
(Ichii and Hada, 1983). In addition, appropriate agronomic management
could largely reduce the lodging risk of the main crop. Decreasing the
N application rate and soil drying during the tillering stage of the main crop,
and adjusting the planting density and spacing significantly were suggested to
significantly reduce the lodging susceptibility of the rice crop (Kashiwagi
et al., 2005; Kim et al., 1995, 1999; Liu et al., 2014). In addition, pre-
harvesting of the main crops immediately after lodging could also mitigate
the yield losses in the ratoon season. However, the application of these
methods should be based on the grain filling degree of the main crops when
lodging occurred. Harvesting the main crops too early would decrease the
grain yield and quality of rice, which would decrease the profits for farmers.

7.2 Mechanical harvesting of main crops


The present commercial rice harvest machine is not suitable for the ratoon
rice system due to the severe rolling damage to the rice stubble, which could
result in great yield losses of the ratoon crops. Our unpublished data revealed
the mechanical harvesting resulted in an 18.7% yield reduction in the ratoon
crops, compared with manual harvesting (Fig. 6), which severely limited the
further extension of a mechanically harvested ratoon rice system. The devel-
opment of specialized harvest machines for ratoon rice is urgently needed to
Ratoon rice technology 159

Fig. 6 The comparison between manual harvesting and mechanical harvesting of the
main crop on the growth performance of the ratoon crop. The photo was shot at the
maturity stage of the ratoon crop.

reduce the yield reduction of ratoon rice under mechanical harvesting. In


recent years, several specialized harvest machines for the ratoon rice system,
such as the head spike harvester (designed by Huazhong Agricultural Uni-
versity, Wuhan, China) and the rear crawler harvester (designed by Huanan
Agricultural University, Guangzhou, China) have been developed. Further
work should focus on the commercial application and extension of these
machines. In addition, several agronomic practices have been recommended
to promote the regeneration of ratoon buds after the mechanical harvesting.
Soil drying during the middle-late grain filling stage of the main crop
increased the soil hardness degree, which could mitigate the rolling damage
to the ratoon crops caused by the harvest machine. Availing the border effect
of the non-rolling zone was another option to compensate the yield loss cau-
sed by mechanical harvesting. It was reported that the border effect of the
ratoon crop from the non-rolling zone could compensate for 4–10% of
the yield losses caused by the rolling damage from the harvest machine
(Xiao, 2017). In addition, adjusting the planting spacing of the main crop
according to the track width of the rice harvester increased the ratoon season
yield by decreasing the area of the rolling zone (unpublished data).

7.3 Variety breeding for ratoon rice


A key component of the success in ratoon rice is the breeding and selection
of suitable cultivars. However, the current studies are mainly focused on the
screening of varieties with high ratooning ability, and no breeding program
has been undertaken to breed specialized varieties for ratoon crops. The ideal
160 Weiqin Wang et al.

varietal traits for rice ratooning include superior ratooning ability (Bollich
and Turner, 1988), good grain quality in both main and ratoon seasons
(Alizadeh and Habibi, 2016), cold tolerance at ratoon flowering (Yang
et al., 2012), high lodging resistance (Ichii and Hada, 1983), medium growth
duration (Lin et al., 2015), high resistance to pests and diseases, etc. In addi-
tion, the ratio between the sink and the source at the heading stage of the
main crops, the stem thickness of the main crops, the tillering ability of the
main crops, and the ratio between the ratooning bud number and total stem
number at 5 days after the harvesting of the main crops can be used as indi-
cators to identify the ratooning ability of the varieties (Xu et al., 2015). In
addition, future research should focus on exploring the inheritance and
genetic mechanisms associated with the ratooning ability to advance the
breeding progress for ratoon rice.

8. Concluding remarks
The present review suggested that ratoon rice is an economic, green
and resource-efficient rice production system with the advantages of
improving the grain quality, increasing the gross returns for farmers, and
decreasing greenhouse gas emissions. Integrated agronomics technologies,
including the selection of high regeneration varieties, the application of
bud-promoting fertilizer and tiller-promoting fertilizer, soil drying during
the main crops season and flooded irrigation immediately after the
harvesting of the main crops, the appropriate stubble cutting height
according to the varietal traits and meteorological conditions, the incorpo-
ration of mechanical harvesting and direct-seeding, and pest and disease pre-
vention are strongly recommended to enhance the productivity and
sustainability of ratoon rice systems. Nevertheless, constraints such as lodg-
ing, lack of specialized rice harvester and the appropriate varieties limit the
wide scale adoption of ratoon rice. Effective strategies to address these con-
straints are needed, including the varieties bred and selected for rice
ratooning, guidelines for enhancing lodging tolerance, and the development
of specialized rice harvest machines for ratoon rice.

Acknowledgments
This work is supported by the National Natural Science Foundation of China (Project No.
31971840) and National Key R&D Program of China (Project No. 2018YFD0301300).
Ratoon rice technology 161

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