B IOD I V E R S I TA S ISSN: 1412-033X

Volume 24, Number 1, January 2023 E-ISSN: 2085-4722

Pages: 208-214 DOI: 10.13057/biodiv/d240125

Genetic diversity, heritability, and accumulation patterns of Zn content

in biofortified rice

WAGE RATNA ROHAENI1,, WILLY BAYUARDI SUWARNO2,, UNTUNG SUSANTO3,

TRIKOSOEMANINGTYAS2, MUNIF GHULAMAHDI2, HAJRIAL ASWIDINNOOR2,
1Graduate Program in Plant Breeding and Biotechnology, Faculty of Agriculture, Institut Pertanian Bogor. Jl. Raya Dramaga, Kampus IPB Dramaga,
Bogor 16680, West Java, Indonesia. Tel./fax.: +62-271-637457 Ext. 129, email: wageratna@apps.ipb.ac.id
2Plant Breeding and Biotechnology Laboratory, Department of Agronomy and Horticulture, Faculty of Agriculture, Institut Pertanian Bogor. Jl. Raya

Dramaga, Kampus IPB Dramaga, Bogor 16680 West Java, Indonesia.  email: hajrial@apps.ipb.ac.id.  willy@apps.ipb.ac.id
3Agricultural Food Research Organization, National Innovation Research Agency. Cibinong Science Center, Jl. Raya Jakarta-Bogor Km. 46, Cibinong,

Bogor 16911, West Java, Indonesia

Manuscript received: 19 October 2022. Revision accepted: 4 January 2023.

Abstract. Rohaeni WR, Suwarno WB, Susanto U, Trikosoemaningtyas, Ghulamahdi M, Aswidinnoor H. 2023. Genetic diversity,
heritability, and accumulation patterns of Zn content in biofortified rice. Biodiversitas 24: 208-214. The study on zinc accumulation
patterns in various rice organs is valuable information to support effective biofortified breeding. Research reports related to zinc
accumulation have only been limited to seeds, while the pattern of zinc accumulation in each part of the plant has not been r eported,
especially for Indonesian rice germplasm. The study aims to provide information on the pattern of accumulation and genetic diversity of
Zn content in rice plants. The experiments used a randomized complete block design (RCBD) with 3 replications and genotypes as a
treatment. The genotypes were 29 lines of BC4F3 Inpari IR Nutri Zinc////Inpari 23, 5 lines of BC4F3 Inpari IR Nutrizinc////Sintanur, and
1 line of BC4F3 Inpari IR Nutrizinc////Inpari 35. A total of 6 varieties were used as checks. The Zn content of dehulled (brown rice)
grains sample was measured using the Oxford Supreme8000 XRF machine with the Rice15 method that had been validated by the
Inductively Coupled Plasma (ICP) method, while the analysis of the pattern of Zn content of several organs (seeds, leaves, stems, and
roots) used the Inductively Coupled Plasma Optical Emission Spectrometry (ICP-IOS) method. The coefficient of genetic variation of
the grain zinc content (GZn) was moderate and had high heritability criteria (H 2bs = 0.83). Yield characters have a narrow coefficient of
genetic variation and low heritability. There were differences in the pattern of Zn accumulation in plant organs between high GZn and
low GZn genotypes. The distribution of Zn content in high GZn genotypes showed 26.4% in seeds, 13.7% in leaves, 39.3% in stems,
and 20.6% in roots. The distribution of Zn content in low GZn genotypes showed 14.2% in seeds, 10.9% in leaves, 31.0% in stems, and
44.0% in roots. The results of this study indicate that a different breeding strategy is needed to develop new high-yielding rice varieties
based on zinc accumulation patterns in cross-parents.

Keywords: Biofortification, genetic diversity, rice, Zn accumulation

INTRODUCTION engineering approaches (biotechnology and conventional

breeding), agronomic approaches (microbe-mediated,
Malnutrition and chronic disorders in developing mineral-rich fertilization, growth regulators, and cultivation
countries are the main obstacles to achieving the systems) (Garg et al. 2018; Olson et al. 2021 ), and post-
Millennium Development Goals (Dipti et al. 2012). harvest product supplementation (Siwela et al. 2020).
Malnutrition, such as stunting, wasting, and being Biofortification through a plant breeding approach is
underweight in childhood, is reported to cause more than reported to be more effective and efficient than agronomy
1.0 million deaths, 3.9% of life years lost, and 3.8% of life and supplementation approaches, especially to reach target
years lost. The prevalence estimation value of malnutrition areas that have limited access to factory-made nutrient-rich
in West African, South Asian, and Southeast Asian products (Olson et al. 2021).
countries are thought to be much higher than the global Biofortification of micronutrients, vitamins, and other
average (Ssentongo et al. 2021). Improving nutrition by important substances, including Zn is possible to be carried
increasing the nutrient content of staple foods, such as rice, out in rice commodities through conventional breeding
will be more effective because it is consumed daily by the approaches (Swamy et al. 2016; Nakandalage et al. 2016;
people of Indonesia. Rice breeding programs can focus on Rao et al. 2020). The effectiveness of selection is largely
developing new varieties with higher Fe, Zn, or β-carotene determined by genetic diversity and the heritability value of
content. Indonesia, one of the developing countries prone the target characters (Keerthana et al. 2019). The higher the
to stunting, has pursued a biofortified rice breeding genetic diversity, the higher the selection effectiveness.
program for zinc to overcome stunting. The selection of characters with moderate to high
Biofortification is defined as an effort to increase the heritability values will be effective in the early generations.
nutritional value of food commodities through genetic Conversely, selection for characters with low heritability
 ROHAENI et al. – Genetic diversity zinc biofortified rice 209

may be more effective in later generations (>F7). 5 lines BC4F3 Inpari IR Nutrizinc////Sintanur, and 1 line
The study of the genetic diversity of Indonesian rice BC4F3 Inpari IR Nutrizinc////Inpari 35. The check varieties
germplasm for the character of zinc content in grain is still used were 6 varieties, namely Inpari IR Nutri Zinc, Inpari
limited. The Indian Institute of Rice Research (IIRR) 23, Sintanur, Inpari 35, Ciherang, and Inpari 46 TDH.
initiated efforts to biofortify rice for Zn by characterizing Research at each location using a randomized complete
the Zn content in various dehulled rice using energy- group design with 3 replications. Seedlings aged 21 hss
dispersive X-ray fluorescence spectroscopy (XRF). The were transplanted with a spacing of 25cm x 25cm in a 4m x
results reported that the zinc content in rice ranged from 5m plot. Fertilizer applications followed a recommendation
7.3 to 52.7 mg/kg (Rao et al. 2020). So far it has been from The experimental station of ICRR (equal to the results
known that Oryza australiensis is a species with high of the paddy soil rapid test kit with a recommendation of
nutritional content. Meanwhile, breeding efforts for Zn-rich 300 kg/ha of urea, 100 kg/ha of SP-36, and 50 kg/ha of
rice in Indonesia have succeeded in obtaining seven lines KCl). Pest and disease management according to Integrated
that have higher grain Zn content (range 30.0-34.2 mg/kg) Crop Protection principles.
than the Ciherang line. These five lines have productivity Several agronomic characteristics that were measured
comparable to Ciherang (Susanto et al. 2017). were plant height (cm), the number of tillers, panicle length
The nutritional content of a grain of rice is influenced (cm), panicle weight per clump (g), total grain weight per
by the morphology and histological characteristics of clump (g), and filled grain weight per clump (g). The
cariopsis (Roy and Shil 2020), both of which are targeted traits were the Zn content in brown rice (mg/kg)
genetically controlled. It is not enough to know the genetic and yield (t/ha at 14% moisture content), and the supported
diversity of zinc in grains, the exploration of genetic biofortified trait was the Fe content in brown rice (mg/kg).
diversity related to Zn content in other organs is also Measurement of zinc content for grains (GZn - Grains
important, because physiologically, zinc is translocated zinc) according to the standard procedure described by
from roots to all plant organs, including rice grains. Rohaeni et al. (2016) and Susanto et al. (2017). The Zn
Therefore, information on the accumulation pattern of Zn content (mg/kg) of the peeled brown rice grain sample was
content and its genetic diversity in various plant organs will measured using the Oxford Supreme 8000 XRF machine
be very useful as a basis for determining future rice with the Rice15 method which has been validated by the
biofortification breeding programs for Zn, both ICP method. The zinc measurement to study accumulation
conventionally and using biotechnology. Estimation by patterns in plant organs was carried out using the
combining heritability values and hereditary propels is Inductively Coupled Plasma Optical Emission
thought to be very helpful in predicting genetic progress Spectrometry (ICP-OES) method at the SIG-Bogor
compared to relying solely on heritability estimation values Company Laboratory. Analysis of the pattern of zinc
(Cooper et al. 2021). Information on genetic diversity and accumulation was carried out on representatives of low and
character control genes related to zinc accumulation in rice high GZn genotypes taken from experimental harvest
is needed to see the effectiveness of selection methods in samples at site 1. A total of 7 genotypes were used for this
biofortified rice breeding programs. The present analysis. Samples of plant organs were taken from 3
investigation aims to provide information on genetic replicates.
diversity and accumulation patterns of Zn content in
several rice organs and to obtain potential lines with high Data analysis
yields and high Zn content in the grain. The analysis covered ANOVA per site, combined
ANOVA, estimation of H2bs value, and coefficient of
genetic variation (CGV) (Table 1). If the genotype effect
MATERIALS AND METHODS was significant at the 5% level, then the least significant
difference (LSD) test at the 5% level was performed.
Study area Statistical analyses were conducted using SAS, Microsoft
The research was conducted at two locations (an Excel, and Minitab.
irrigation agroecosystem and rainfed lowland) in March -
September 2022. The site 1 irrigation agroecosystem was
located at The experimental station of the Indonesian
Center of Rice Research (ICRR), Subang, West Java - Table 1. Combined analysis and partition of expectation of mean
Indonesia (6°21'10.0"S:107°38'54.1"E with an altitude of square for estimating genetic variation component
10m above sea level, the soil type was Alluvial). The site 2
rainfed lowland was located at 6°31'08.4"S:107°31'05.3"E Source DF MS E(MS)
with an altitude of 92.7m above sea level and the soil type Environment (E) l-1 M5
was Latosol. Replication/environment l(r-1) M4
Genotype (G) g-1 M3 σ2e+r σ2gl + rl σ2g
GxE (g-1) (l-1) M2 σ2e + r σ2gl
Genetic diversity and heritability of agronomic
Error (E) l(g-1)(r-1) M1 σ2e
characters and Zn content Notes: DF: degree of freedom, MS: mean square, E(MS):
A total of 35 promising lines were used in this study, expectation of mean square
including 29 lines BC4F3 Inpari IR Nutri Zinc////Inpari 23,
210 B I OD I V E R S ITA S 24 (1): 208-214, January 2023

Estimation of the components of genetic, The heritability value with high criteria was owned by
environmental, phenotype, and GxE variation were referred GZn (H2bs = 0.83), 1000 grain weight (H2bs = 0.91), and
to in Table 1 and carried out in the following formulas: maturity age (H2bs = 0.65). While the heritability value with
σ2 G = (M3 - M2) / rl moderate criteria was on plant height, number of tillers,
σ2 GxE = (M2 - M1) / r number of panicles per clump, number of filled grain per
σ2 E = M1 clump, number of empty grains per clump, and grain Fe
σ2 P = σ2 G + σ2 E / rl + σ2 GxE / l content (Table 2). These results led to the notion that the
Where: l: environment, r: replication, σ2 E: environment GZn trait, which was the target of the study, had a very
variance, σ2 G: genotype variance, σ2 P: phenotype large potential to be inherited, indicated by a high criterion
variance, σ2 GxE = GxE variance heritability value. This result is the same as several previous
reports showing that GZn has high heritability (Swamy et
al. 2016; Velu et al. 2019; Mukamuhirwa et al. 2015).
The GZn character, being the target character of this
study, shows a high heritability value. This means that the
criteria :
selection of high Zn lines can be carried out from the early
Low (Hbs2 ≤ 20%); Moderate (20% < Hbs2 < 50%); High
generation. Traits with high heritability predictive value
(Hbs2 ≥ 50%).
have higher prospects for selection success because most of
the variation is caused by genetics. Estimates of heritability
Coefficient of genetic variation (Singh and Chaudhary
provide a better estimate of population genetic diversity
2007):
and allow prediction of selection gain. Therefore, they are
very important in breeding programs (Ribeiro et al. 2019).
Sadimantara et al. (2021) reported that a high heritability
Where: predictive value helps select superior genotypes based on
= general mean phenotypic appearance. It has been reported that rice has
CGV categories: narrow (0-10%), moderate (10-20%), abundant germplasm variation thus providing the required
and wide (> 20%) genetic diversity for rice breeding programs. The existence
of genetic variability is a prerequisite for a successful
breeding program for any species. Sadimantara et al.
RESULTS AND DISCUSSION (2021) showed that the yield character has a heritability
value with high criteria and a moderate coefficient of
Genetic diversity and heritability of agronomic genetic variation.
characters and Zn content The results of this study indicated that there were
The estimated value of genetic diversity showed the considerable challenges in combining the yield character
character of plant height, number of tillers, physiological with the GZn character. The heritability value of the GZn
maturity, number of panicles per hill, number of filled was high, while the yield character was low. The selection
grains per hill, number of empty grains per hill, 1000 grain for high GZn can be performed in early generations, but
weight, and grain Zn content (GZn) had a coefficient of selection for high yield will perhaps be more effective
genetic variation with moderate criteria. Meanwhile, the when carried out in the later generations. GZn selection in
maturity age, the weight of filled grain per clump, the Fe the early generations might allow accumulating the
content of the grain, and the yield characters had narrow additive genes regulating GZn, but there will be erosions of
genetic diversity (CGV) (Table 2). additive genes controlling yield.

Table 2. Estimated variance components and broad-sense heritability of biofortified rice genotypes based on field trials from two
agroecosystems (irrigation and rainfed lowland)

Characters Mean CV % σ2 g σ2gxe σ2 e σ2 p CGV % CGV criteria H2bs H2bs criteria

PH 105.72 11.85 147.69 2.10 157.07 306.86 11.50 Moderate 0.48 Moderate
NT 18.02 16.19 5.21 0.50 8.52 14.23 12.66 Moderate 0.37 Moderate
MA 113.32 1.20 5.34 1.04 1.86 8.24 2.04 Narrow 0.65 High
NP 18.02 16.19 5.21 0.50 8.52 14.23 12.66 Moderate 0.37 Moderate
NPG 1928.32 15.76 50065.88 3190.44 92351.59 145607.91 11.60 Moderate 0.34 Moderate
NEG 404.49 24.16 2411.04 217.43 9552.56 12181.03 12.14 Moderate 0.2 Moderate
WPG 44.31 16.6 5.85 0.81 54.14 49.10 5.46 Narrow 0.12 Low
1000GW 23.25 3.66 7.96 0.02 0.72 8.70 12.13 Moderate 0.91 High
FE 16.54 9.50 1.31 0.05 2.47 3.82 6.91 Narrow 0.34 Moderate
ZN 29.45 6.20 23.95 1.44 3.33 28.73 16.62 Moderate 0.83 High
Yield 6.67 12.30 0.13 0.16 0.67 0.96 5.41 Narrow 0.13 Low
Notes: PH: plant height (cm), NT: number of tillers, MA: maturity age, NP: number of panicles per clump, NPG: number of filled grains
per clump, NEG: number of empty grains per clump, WPG: weight of filled grains per clump, 1000GW: 1000 grains weight, FE: grains
Fe content (mg/kg), ZN: grains Zn content (mg/kg), CV: coefficient of variation (experimental error), CGV: coefficient of genetic
variation, H2bs: Broad-sense heritability
 ROHAENI et al. – Genetic diversity zinc biofortified rice 211

Table 3. The best eight genotypes based on higher GZn than the best check (Inpari IR Nutri Zinc)

GEN PH NT MA NP NPG NEG WPG 1000GW FE ZN Yield

G17 104.47 18.85 111.83 17.33 1771.56 392.61 42.17 22.97 19.07+ 33.07+ 6.66
G32 105.46 21.22+ 112.80 17.20 1912.40 463.00 43.39 22.98 16.60 32.04 6.88
G13 101.38 18.43 111.50 18.11 1851.17 388.44 44.54 24.18+ 16.33 31.65 7.17
G5 99.57 19.72 111.50 18.61 1772.39 392.72 41.87 23.80+ 15.75 31.02 6.76
G11 108.77+ 19.28 114.33+ 17.61 1869.72 384.22 42.62 22.93 16.28 30.87 6.71
G20 104.40 18.07 113.50 16.28 1605.50 385.94 38.23 23.77+ 16.62 30.82 6.96
G4 104.82 18.55 113.00 18.11 2184.83+ 316.94 48.00 22.20 16.30 30.10 7.00
G6 101.63 17.75 113.00 18.72 2024.11+ 436.44 45.19 22.38 16.13 30.02 7.46
Inpari IR Nutri Zinc 103.68 19.15 112.50 18.33 2183.44+ 391.22 47.99 22.38 16.62 29.98 6.61
Mean 105.72 19.02 113.32 18.02 1928.33 404.48 44.34 23.25 16.54 29.45 6.67
CV % 11.85 13.75 1.20 16.23 15.76 24.16 16.57 3.66 9.50 6.20 12.30
LSD 5% 2.99 0.94 1.56 3.34 347.01 111.61 8.39 0.97 1.79 2.08 0.94
Notes: PH: plant height (cm), NT: number of tillers, MD: maturity age, NP: number of panicles per clump, NPG: number of filled grains
per clump, NEG: number of empty grains per clump, WPG: weight of pithy grains per clump, 1000GW: 1000 grains weight, FE: grains
Fe content (mg/kg), ZN: grains Zn content (mg/kg), Yield: ton/ha. +: significantly higher than check (Inpari IR Nutri Zinc)

Several strategies are needed to combine 2 traits that A breeding strategy involving crossing combinations is
have different heritability criteria (low and high) such as needed to increase the chances of obtaining high GZn
GZn and yield. First, do pre-breeding for trait enrichment genotypes as well as high yields. Breeding high-yield
in prospective crossbreeding parents. Second, increasing varieties that are responsive to zinc-rich fertilization can
the number of populations and various types of crosses. increase the GZn content of high-yield genotypes.
Third, using a new plant breeding technique approach by Muthukumararaja and Sriramachandrasekharan (2012)
inserting the GZn control gene from super high zinc showed that rice plants are very responsive to the addition
parents into the genome of high-yielding varieties with of extracellular zinc. Shahzad et al. (2021) reported that
high yield potential. plant biofortification could be achieved through
Fortunately, this study resulted in some new promising conventional approaches or new breeding techniques
lines with superior GZn content and higher yield potential (NBTs). Conventional approaches to biofortification
than the check variety. The G17, G32, G13, G5, G11, G20, include mineral fertilization through foliar or soil
G4, and G6 were the promising lines that had higher GZn applications (Cakmak and Kutman 2018), as well as
than the best GZn parents (Inpari IR Nutri Zinc, GZn 29.98 microbial-mediated nutrient uptake enhancement (Krithika
mg/kg) and had better yield potential (Table 3). Lines G6 and Balachandar 2016).
(GZn = 30.02 mg/kg; Yield = 7.46 tons/ha), G13 (GZn =
31.65 mg/kg; Yield = 7.17 tons/ha), and G4 (GZn = 30.10 Zinc distribution pattern on rice plant organs
mg/kg; Yield = 7.00 tons/ ha) showed yield potential with Analysis of variance showed that replication had no
better GZn content than the high GZn parent Inpari IR significant effect on Zn content in all rice plant organs.
Nutri Zinc (GZn 29.98 mg/kg; Yield = 6.61 mg/kg). Genotype has a very significant effect on the diversity of
Furthermore, these promising lines need to be tested for zinc content in each organ (Table 4). Further tests in this
further yields in several locations to see the stability of the study showed that there were 3 classes of genotypes based
GZn content and potential yields (Table 3). on GZn content. The classes were genotypes containing
Conventional breeding for the combination of high, moderate, and low GZn. The high GZn was possessed
biofortified and high-yield rice traits requires more energy by the Pulut Timuru genotype (34.0 mg/kg) and was
for several reasons. First, the correlation between the GZn significantly different from the moderate and low
trait and yield properties is inconsistent. Some reports show genotypes. Genotypes included in the moderate GZn
a negative correlation between grain zinc trait and yield category were Inpari IR Nutri Zinc (19.6 mg/kg), G5 (21.2
components in rice (Inabangan-Asilo et al. 2019; Rao et al. mg/kg), Inpari 23 (18.2 mg/kg), and G27 (18.4 mg/kg). The
2020) and wheat (Khokhar et al. 2020), but several other genotypes included in the low GZn category were Inpari 46
studies show the opposite results (Paramesh et al. 2020; GSR TDH (16.5 mg/kg) and G34 (13.9 mg/kg) (Table 5).
Calayugan et al. 2020). Second, selection for high GZn The present study showed that there were differences in
may be effective in early generations, whereas selection for the pattern of Zn accumulation in plant organs between
a high-yield trait is effective in later generations. GZn has high GZn and low-moderate GZn genotypes. The
high heritability but the yielding character has low distribution of Zn content in the high GZn genotype, Pulut
heritability. Third, in the present study, the genetic Timuru, showed higher content of zinc (26.4%) in seeds
diversity for GZn was included in the moderate criteria but than in stems (39.3%) while zinc content in leaves was
the genetic diversity for the yield components was included 13.7%, and 20.6% in roots.
in the low category.
212 B I OD I V E R S ITA S 24 (1): 208-214, January 2023

Table 4. Analysis of variance on the character of Zn content on seeds, leaves, stems, and roots

F value
Source of variation
Zn grain Zn leaf Zn stem Zn root Zn total
Replication 0.07 ns 0.72 ns 2.81 ns 0.42 ns 1.85 ns
Genotype 33.64 ** 12.89 ** 10.97 ** 3.67 * 4.68 *
Note: *, **: significant at p < 0.05 and p < 0.01, respectively; ns: not significant

Table 5. Zn content (mg/kg) in grain, leaf, stem, and root on low, moderate, and high GZn genotype

Genotype Zn grain Zn leaf Zn stem Zn root Zn total

High GZn genotype
Pulut Timuru 34.0a 17.7a 50.6a 26.6c 128.9a
Moderate GZn genotypes
Inpari IR Nutri Zinc 19.6bc 14.6b 18.4c 38.0bc 90.6c
G5 21.2b 13.5bc 20.5c 50.8ab 106.0bc
Inpari 23 18.2bc 14.0b 24.7bc 37.1bc 94.0c
bc b bc
G27 18.4 14.3 24.1 42.1ab 98.9bc
Low GZn genotypes
Inpari 46 TDH GSR 16.5cd 11.4d 32.0b 39.5bc 99.4bc
G34 13.9d 11.8cd 34.2b 54.5a 114.4ab
Note: Means followed by the same letter in the same column are not significantly different. GZn: grain zinc content (mg/kg)

approach can be applied to genotypes with accumulation

patterns, such as G34 (Table 5). Meanwhile, zinc
accumulation in Pulut Timuru, a zinc-rich genotype, was
concentrated in the stems, whereas in the moderate and low
GZn genotypes, zinc accumulation was generally found in
the root organs, as reported by Impa et al. (2013) and
Stomph et al. (2014).
Rice is reported to be capable of accumulating large
amounts of heavy metals in its seeds (Uddin et al. 2021).
This accumulator property is used to enrich the nutritional
content of rice with useful minerals. However, knowledge
of the physiological processes in mineral absorption needs
to be considered. In rice, zinc is absorbed by the roots and
can be mobilized to all parts of the plant (Impa et al. 2013;
Figure 1. Percentage of zinc content in grains, leaves, stems, and Stomph et al. 2014). In contrast, iron is local, stored in
roots in high, moderate, and low GZn genotypes certain cellular components, and has a certain physiological
role. Post-anthesis Zn uptake by shoots increased and Zn
translocation from roots to shoots decreased as the Zn
Conserve, the distribution pattern of Zn content in low application rate increased through fertilization. Zn
GZn genotype, such G34 and Inpari 46 TDH, showed accumulation rate in grain resulting from pre-anthesis vs.
14.2% in seeds, less than zinc accumulation in stem post-anthesis remobilization depends on the availability of
(31.0%), while zinc accumulation on leaves and root, were Zn in the soil. Pre-anthesis remobilization occurs at lower
10.9% and 44.0%, respectively. The unique pattern of zinc soil Zn content (Liu et al. 2019).
accumulation was found in the moderate GZn genotype. The OsZIP1 and OsZIP3 genes have been shown to be
The first pattern was followed high GZn genotype (20.8% associated with Zn uptake in rice roots and Zn homeostasis
in seeds, 19.8% in stems, 14.3% in leaves, and 45.2% in in rice shoots, whereas OsZIP4 transports Zn intracellularly
roots), called genotype type I, and the second pattern was or between rice tissues (Ishimaru et al. 2007). The
according to a low GZn genotype (19.0% in seeds, 25.3% mechanism of Zn accumulation in each rice organ can be
in stems, 14.7% in leaves, and 41.1% in roots), namely grouped into two categories: Zn is absorbed and retained in
genotype type II (Figure 1). the root tissue, or remobilization of Zn from root to shoot
The G34 line was the lowest GZn, but the total zinc (Impa et al. 2013). Genotypes with concentrated Zn
value in all organs (114.4 mg/kg) was equivalent to the accumulation patterns in roots require a genetic
highest GZn genotype (Pulut Timuru = 128.9 mg/kg). This engineering approach to increase Zn accumulation in seeds.
indicates that G34 line can be increased in GZn content by On the other hand, genotypes capable of accumulating zinc
genetic engineering to relocate the zinc content from the up to the stem are also indicated to be able to translocate
stem to the seed. The gene overexpression technology high amounts of zinc to the seed organs, such as in Pulut
 ROHAENI et al. – Genetic diversity zinc biofortified rice 213

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