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Biotechnology for Jatropha improvement: A worthy exploration

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 Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277

Contents lists available at ScienceDirect

Renewable and Sustainable Energy Reviews

journal homepage: www.elsevier.com/locate/rser

Biotechnology for Jatropha improvement: A worthy exploration

M. Moniruzzaman a, Zahira Yaakob a,n, Rahima Khatun a,b
a
Department of Chemical and Process Engineering, Faculty of Engineering and Built Environment, Universiti Kebangsaan Malaysia, Bangi 43600, Malaysia
b
Department of Secondary and Higher Education, Ministry of Education, Bangladesh

art ic l e i nf o a b s t r a c t

Article history: Comfortable life, economic growth, industrialisation, global warming, energy security and sustainable
Received 28 April 2015 environment are burning issues facing modern civilisation. The availability of adequate renewable energy
Received in revised form is in demand. Jatropha is being explored as a potential biofuel crop candidate because of its biodiesel
20 September 2015
production potential, high oil content, rapid growth, easy propagation, drought tolerant nature, relatively
Accepted 23 October 2015
less irrigation and agricultural inputs, insect and pest resistance. However, previous programmes for
Jatropha plantation did not satisfy the expectation because of the absence of a good commercial variety,
Keywords: large scale propagation without evaluating the planting material, knowledge gap and consideration as
Jatropha improvement low a impute crop. Lack of systematic breeding programmes, the inexistence of a collaboration between
Biotechnology
scientists in this field, the unavailability of desired germplasm and more importantly less variability
Cutting edge
within the species are the constraints for the conventional breeding for a Jatropha improvement pro-
Renewable energy
gramme. Biological techniques have proven records for the improvement of many crops. Jatropha
“organogenesis”, which has insignificant contribution to genetic improvement, is studied. Several
genomic and transgenic studies have been reported, but it is still far behind in comparison to other crops.
It is time to investigate somaclonal variation, in vitro selection and haploid breeding for Jatropha
improvement. Resequencing and transcriptom analysis are necessary for high dance linkage map and a
good reference genome. Genome wide association studies (GWAS) and genomic selection (GS) are
pending. Genetic engineering, particularly to increase female flowers in inflorescence, eliminates the
toxic component and increases tolerance to diseases, insects and pests should be given priority.
& 2015 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1263
1.1. Limitation of Jatropha as a biofuel crop . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1263
1.2. Jatropha breeding objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1263
2. Potentials of Jatropha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1264
2.1. Jatropha as energy source . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1264
2.2. Industrial use . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1265
2.3. Other uses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1265
3. Conventional breeding of Jatropha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1265
3.1. Limitation of conventional breeding for Jatropha improvement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1266
4. Biotechnology for Jatropha improvement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1266
4.1. Tissue culture. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1268
4.2. Organogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1268
4.3. Haploid plant production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1270
4.4. Embryo culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1270
4.5. Endosperm culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1270
4.6. Somatic hybridisation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1270
4.7. Mutagenesis in crop improvement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1271
4.8. Somaclonal variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1271

n
Corresponding author. Tel.: þ 60 389216420; fax: þ 60 389216148.
E-mail address: zahirayaakob65@gmail.com (Z. Yaakob).

http://dx.doi.org/10.1016/j.rser.2015.10.074
1364-0321/& 2015 Elsevier Ltd. All rights reserved.
 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277 1263

4.9. Genomic resources- DNA markers, marker assisted selection and molecular breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1272
4.10. Genetic engineering. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1273
5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1273
Acknowledgement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1274
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1274

1. Introduction Jatropha biomass as renewable material for biocomposites [25]; and

the status of Jatropha cultivation in major cultivating countries, such
Jatropha belongs to the family Euphorbiaceae and has 175 as China [26–28], India [29], Malaysia [30,31] and Indonesia [32].
species. It originated from tropical America and distributes allover Focusing on potential but unexplored areas of biotechniques as well
the tropics and subtropics of Asia and Africa [1]. These plant as studied areas of biotechniques for Jatropha improvement makes
species are being used as hedges, wind protection, soil erosion this study distinct from previous studies.
control, firewood and some sort of therapeutic agent by tribal
communities [2–5]. The oil from the seeds is also being used for 1.1. Limitation of Jatropha as a biofuel crop
lamp fuel, soap making, paints and as a lubricant [3,6,7].
The world energy spending has been increasing, and during the
period of 50 years ending in 2025, it will have increased three fold. 1. A high productive commercial variety has not yet been
Over 80% of the total combusted energy is still coming from fossil developed.
fuel. The fossil fuel oil is expected to face a greater threat of 2. The trees vary greatly in yield.
becoming exhausted. Global oil production may likely reach a 3. It can survive on insufficient irrigation and nutrients, but the
maximum between 2015 and 2030. It is then expected to sharply production is reduced.
decline to 40% of today's production by 2075 [8]. 4. 3–5 years are required for commercial productivity, which is
The plant “Jatropha” is an important crop because of its bio- longer than annual oilseed crops.
diesel production potential, high oil content, rapid growth, easy 5. The presence of toxins prevents the use of the seed cake for
propagation, drought tolerant nature, ability to grow and reclaim livestock feed.
various types of land, need for less irrigation and less agricultural 6. Pests and diseases susceptibility of Jatropha.
inputs, pest resistance, short gestation periods and suitable traits 7. Jatropha is affected by frost and water logging.
for easy harvesting [9,10]. In addition, this crop is insect and pest 8. Jatropha may act as a host for cassava diseases.
tolerant, can grow on degraded and marginal lands and animals do 9. The high viscosity of Jatropha oil is not suitable for cool climate
not graze on it [11]. Biofuel production from edible crops (e.g., conditions.
corn, soya, and palm) competes with global food security. Using 10. Jatropha may become a weed problem in certain environments
Jatropha, a non-food crop, as renewable energy crop can mitigate [11,33]:
this problem [12,13].
There are a number of good reviews (Table 1) on Jatropha. Some 1.2. Jatropha breeding objectives
of them address general, socio-economic, environmental and sus-
tainability aspects of Jatropha cultivation [14–19]; comparative
studies on the performance evaluation of Jatropha biodiesel [20– 1. Improve the dry matter distribution with greater assignment to
22]; biotechnological approaches for improving Jatropha [10,23,24]; fruit rather than vegetative parts.

Table 1
Review articles on Jatropha.

1 Jatropha curcas L.: a crucified plant waiting for resurgence. [10]

2 Comparative studies on performance evaluation of DI diesel engine with high grade low heat rejection combustion chamber with carbureted alcohols and [21]
crude Jatropha oil.
3 Performance evaluation of medium grade low heat rejection diesel engine with carbureted methanol and crude Jatropha oil. [22]
4 Global experience with Jatropha cultivation for bioenergy: an assessment of socio-economic and environmental aspects. [15]
5 Jatropha curcas:: a ten year story from hope to despair. [163]
6 Jatropha curcas as a renewable source for bio-fuels—a review. [16]
7 A Jatropha biomass as renewable materials for biocomposites and its applications. [25]
8 Status of molecular breeding for improving Jatropha curcas and biodiesel. [164]
9 A global comparative review of biodiesel production from Jatropha curcas using different homogeneous acid and alkaline catalysts: study of physical and [17]
chemical properties.
10 Land availability of Jatropha production in Malaysia. [30]
11 Prospects of biodiesel from Jatropha in Malaysia. [31]
12 Jatropha curcas: a potential biofuel plant for sustainable environmental development. [14]
13 Review and prospects of Jatropha biodiesel industry in China. [26]
14 Risk management for Jatropha curcas based biodiesel industry of Panzhihua Prefecture in Southwest China. [27]
15 Sustainability issues for promotion of Jatropha biodiesel in Indian scenario: a review. [165]
16 A review of biodiesel production from Jatropha curcas L. oil. [166]
17 Comparison of palm oil, Jatropha curcas and Calophyllum inophyllum for biodiesel: a review. [167]
18 Life cycle assessment of biodiesel from soybean, Jatropha and microalgae in China conditions. [28]
19 Applications of biotechnology and biochemical engineering for the improvement of Jatropha and biodiesel: a review. [24]
20 A review on prospect of Jatropha curcas for biodiesel in Indonesia. [32]
21 Biodiesel production from Jatropha curcas oil. [168]
22 Prospects of biodiesel from Jatropha in India: a review. [29]
23 Biology and genetic improvement of Jatropha curcas L.: a review. [33]
1264 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
2. Increase the ration of female flowers to male flowers per
inflorescence to improve yield.
3. Increase synchronisation of flowering and fruit maturity to
reduce labour requirements and introduce mechanisation of
harvesting.
4. Increase seed weight and seed oil content.
5. Increase the number of branches, flowers, fruits and seeds.
6. Improve the oil quality.
7. Develop a non-toxic variety to ensure human safety and add
value by using the seed cake as fodder.
8. Improve the drought resistance and productivity under stress
conditions.
9. Develop disease and pest tolerant varieties to avoid the use of
costly agrochemicals.
10. Improve plant architecture for dipper rooting and smaller.
2. Potentials of Jatropha
The small plant Jatropha has many uses (Fig. 1). Oil from Jatropha
is an excellent source of renewable energy in biodiesel foams and jet
fuel. The biomass (wood, leaves, and fruits) from Jatropha is com-
bustible and is used as fire wood in rural areas. Jatropha also has
industrial uses. Jatropha oil is used in soap making, cosmetic and
dyeing industries as dye for cloth, and fishing nets. Traditionally,
Jatropha has been used as a medicinal plant. Jatropha possesses
therapeutic compounds that can be used in the modern pharma-
ceutical industry for antimicrobial, anti-inflammatory, healing,
Fig. 1. Uses and energy values of different parts of Jatropha curcas. A energy values of the components are given in MJ kg  1.
Source: [212–214].
homoeostatic, anticholinesterase, antidiarrheal, antihypertensive
and anticancer agents. Toxicological studies are important before the
use of Jatropha and/or its derivatives as a therapeutic agent. The
seed cake of Jatropha contains a high percentage of protein and
other nutrients that can be used as animal feed and organic fertiliser.
Hedges and erosion control are ancient uses of Jatropha.
2.1. Jatropha as energy source
Biomass has been receiving attention as an alternative to fossil
fuel and a source of renewable energy. Production ease, sustain-
ability and environmental friendly properties make biomass
lucrative. A number of crops have been identified and being grown
in energy crop farming as feedstocks for first generation biofuels.
Among these energy crops, Jatropha curcas L. (JCL) has been eval-
uated as the most competent energy crop in tropical regions [34].
Wood, fruit shells, seed husks and kernels are the components
of Jatropha from which energy is derived [35]. Raw oil is the major
energy source from Jatropha. The decorticated seeds contain 40–
60% oil, depending on the variety [36–41]. The oil is being utilised
for many purposes, such as lighting, as a lubricant, for making soap
[42] and most importantly as biodiesel. It satisfies the biodiesel
properties of the European standards (Table 2).
Jatropha oil contains crude protein, crude fat and moisture of
approximately 24.60%, 47.25%, and 5.54%, respectively [43]. The oil
contains both saturated and unsaturated fatty acids. Palmitic acid
(16:0) at 14.1% and stearic acid (18:0) at 6.7% are the major satu-
rated fatty acids, and oleic acid (18:1) at 47.0% and linoleic acid
 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
Table 2
Characteristics of Jatropha biodiesel compared to European specifications: [169].
Characteristic Jatropha European
biodiesel standard
Density (g cm-3 at 20 °C) 0.87 0.86–0.90
Flash point (°C) 191 4101
Cetane no. (ISO 5165) 57–62 451
Viscosity mm²/s at 40 ºC 4.20 3.5–5.0
Net cal. val. (MJ/L) 34.4 – –
Iodine no. 95–106 o 120
Sulphated ash 0.014 o 0.02
Carbon residue 0.025 o 0.3
a
þ indicates that Jatropha performs better than the European standard.
(18:2) at 31.6% are the major unsaturated fatty acids. The useful-
ness of Jatropha oil and its esters instead of petrodiesel has been
reported [44]. The energy value (39 MJ kg  1) of Jatropha seed oil is
higher than anthracite coal and is comparable to crude oil [45].
The shells possess approximately 35–40% of dry fruit, which is
60–65% seed by weight [46]. The seed comprises approximately
42% husks and 58% kernel [35]. The gross energy value
(24 MJ kg  1) of the seed is higher than lignite coal and cattle
manure and is comparable to corn cobs [47].
Mechanical removal of the shell from the fruit is the first step of
oil extraction. The biomolecules cellulose (34%), hemicelluloses
(10%) and lignin (12%) are the chemical constituents of Jatropha
shell [35]. One (1) kg seed shell of Jatropha yields 11.1 MJ energy
[45].The seed husk is made of ash (4%), volatile matter (71%) and
fixed carbon (25%). The energy generated from the seed husk is
comparable to wood. One(1) kg seed husks generates 16 MJ energy
[46].
Hydrogen (H2) gas became very attractive renewable energy for
future because of its less energy expenditures and the possibility
to use low cost organic wastes as substrate. De-oiled Jatropha solid
waste (DJSW) and Jatropha seed cake contain Lignocellulose that
can be used as substrates for fermentation to produce Lig-
nocellulose biohydrogen [48–51]. Kuma et al. studied hydrogen
fermentation process from de-oiled Jatropha waste. It was
observed that at optimum condition, substrate concentration
211 g/L, pH- 6.5 and temperature- 55.4 °C, the highest achiev-
able cumulative hydrogen production (CHP) was 296 mL H2 [50].
Lopes et al. studied dark fermentation of seed cake by a pure strain
of the bacteria Enterobacter aerogenes. They produced 68.2 mL H2/
gVSiJSC biohydrogen without pretreatment of substrate. It is sig-
nificant in the viewpoint of energy saving. They found that, the
increase concentration press cake from 2.5 to 10 gVS/LFM led to the
increase of the cumulative hydrogen production and to higher bio
H2 production [51].
2.2. Industrial use
The viscous oil from Jatropha fruit can be used for soap making
[41]. The high level of palmitic acid and the hydrophobicity of
Jatropha oil make the manufacturing of soft and durable soap easy.
It is being used in West Africa, Zambia, Tanzania and Zimbabwe as
a soapstock, including in the manufacture of soft laundry soap
[52]. Jatropha oil gives a very good foaming, white soap with
positive effects on the skin, partly due to the glycerine content of
the soap [53]. Jatropha soap is said to have medicinal character-
istics and is therefore used by people with various skin diseases
and sensitivity to regular soap [54]. The 36% linoleic acid (C18:2)
content in Jatropha kernel oil is of possible interest for skin care
[52,55]. The oil is also an ingredient in hair conditioners [11].
Many phytochemical constituents, alkaloids, coumarins, flavo-
noids, lignoids, phenols, saponins, steroids, tannins, and terpenoids,
1265
were previously detected in different extracts from different parts of
this plant [56]. Pharmacological studies have demonstrated significant
action of different extracts and/or isolated compounds as anti-
Remarksa
microbial [57], anti-inflammatory [58–60], healing, homoeostatic [61],
anticholinesterase [62,63], antidiarrheal [64–66], antihypertensive
þ [67], and anticancer agents [68,69]. Toxicological studies associated
þþ with phytochemical analysis are important to understand the even-
þþþ
þ
tual toxic effects that could reduce its medicinal value.
The oil has a strong purgative action and is widely used for skin
þ diseases and to soothe pain such as that caused by rheumatism.
þ Due to its antimicrobial properties, the plant is being used in
þþ
pesticides. The bark of Jatropha yields blue dye, which is used for
colouring cloths, finishing nets and lines in the Philippines [70].
The seed oil can be applied to treat eczema [9]. The oil is also an
ingredient in hair conditioners [11].The use of Jatropha for the
soap industry (as an alternative to Karitee butter), and cosmetics
are regarded as one of its non-energy uses.
2.3. Other uses
One of the ancient uses of Jatropha is use as a hedge. The
advantage is that animals do not feed on it. Plantings from ger-
minating seeds produce taproots as well as surface roots; thus, it is
used as a barrier to soil erosion and a nutrient pump because they
are able to extract minerals that have leached down through the
soil profile and return them to the surface through leaf fall, fruit
debris and other organic remains.
Jatropha seed cake is high in protein, 58.1 percent (%) by
weight, compared to soy meal’s 48%, and it would be a valuable
livestock protein feed supplement if it were not for its toxicity.
Nitrogen, phosphorus, potassium, calcium, magnesium, sulphur,
iron, zinc, copper, and manganese are the available nutrients in the
plant, which makes Jatropha seed cake an excellent organic ferti-
liser [71,72].
3. Conventional breeding of Jatropha
Plant breeding is the application and practice of the scientific
principle of altering the genetic pattern of plants to increase their
agronomic and economic value. The main objectives are higher
yields, improved quality, disease and insect resistance, change in
maturity duration, agronomic character (i.e., plant height, tillering,
and branching), photosensitivity, synchronous maturity, non-
shattering characteristics, determinate growth, dormancy, creation
of new varieties for different seasons, moisture stress and salt
tolerance, and elimination of toxic substances. The specific
objectives vary greatly depending on the crop under consideration.
The main activity (Fig. 2) of plant breeding includes i) creation of
variation, ii) selection, ii) evaluation, iii) multiplication and iv)
distribution [73]. Genetic variation can be created by domestica-
tion, germplasm collection, plant introduction, cross pollination
(hybridisation) and biotechnologically by inter-species hybridisa-
tion, mutation, polyploidy, somaclonal variation, germaclonal
variation, and genetic engineering. Searching, domestication,
categorisation and assessment of global germplasm will offer a
solid foundation for developing suitable varieties by different
methods. Wageningen University and Research Centre, Plant
Research International, the Netherlands, Lao Institute of Renew-
able Energy (LIRE), Surfactant and Bio-energy Resource Centre
(SBRC) in Bogor, the Centro Hispaniola de Investigación en Bioe-
nergias y Agricultura Sostenible (CHIBAS) – binational (Haiti and
Dominican Republic), and the Agricultural Research Trust (ART),
Zimbabwe are good sources of Jatropha germplasm.
Selection is the rate limiting step of a successful breeding
programme. Selection is based on the objective of the particular
1266 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
Fig. 2. Activities of plant breeding.
breeding programme. Conventionally, selection is based on phe-
notypic data. Selection breeding involves selection of candidate
plants based on desired traits (Fig. 3). There are some authentic
and short term selection techniques based on biotechnology, such
as marker assisted selection (MAS) and genomic selection (GS).
The term “Evaluation” in breeding is the checking of the perfor-
mance of emerging varieties under actual field conditions. If the
new variety performs well, then it undergoes multiplication and
distribution.
3.1. Limitation of conventional breeding for Jatropha improvement
The shortcomings of conventional breeding are i) it is laborious
and time consuming. Several years are needed to develop a variety,
ii) selection for desired traits may not be accurate, iii) wild crosses
may not be possible due to species boundaries. Inclusive work on
aggregation, categorisation, and assessment of the germplasm for
growth, production, agronomy, and morphology is still immature
for Jatropha breeding.
Molecular study reveals that the genetic diversity among
Jatropha is low [74–79]. Accessions from Mexico and Costa Rica
had a 70% similarity to accessions from other parts of the world
[80]. This may minimise the potential of conventional cross-
pollinated breeding programmes for Jatropha curcas. Marker
assisted selection (MAS) and genomic selection (GS) are authentic
selections of desired traits. They drastically reduce the generation
interval so a desired novel variety can be produced within a short
time. These techniques are well-adapted for many breeding pro-
grammes for both plants and animals. However, the application
and adaptation of these potential techniques are still lacking for
Jatropha breeding.
4. Biotechnology for Jatropha improvement
Agricultural biotechnology is the branch of biotechnology that
addresses plants, animals and microbes to enhance their value.
Genetic improvement of desired crops is one of the basic targets of
agricultural biotechnology. Genetic variability of desired traits is a
prerequisite to become successful in any genetic improvement
programme [81].
 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277 1267

The narrow genetic diversity of Jatropha [74–79] minimises
the potential of intra-specific breeding programmes for Jatropha
curcas. The same studies also raised the prospect of biotechno-
logical intervention (Table 3)- inter-specific hybridisation, hap-
loid production, somatic hybridisation, somatic embryogenesis,
somaclonal variation, mutation breeding, marker assisted
selection and molecular breeding, genetic engineering and
transgenesis. Scientists have to evaluate and develop tools that
are more suitable and fruitful for generating the wanted geno-
types [82].

Fig. 3. Selection breeding approach of Jatropha curcas L.

Table 3
Biotechnological intervention for crop improvement.

Technique Target

Micropropagation Clonal maintenance. Large-scale multiplication, hybrid seed production

Embryo rescue Wide crosses, widening genetic base. transfer of novel traits
Somatic embryogenesis Mutation induction, somaclonal variation, selection, artificial seeds
In vitro fertilisation Use the of self-incompatibility in hybrid variety production
Anther/ovary culture Doubled haploid, inbred lines for hybrids, synthetic, composites
Protoplast fusion Widening genetic base, gene reansfer
Organelle transfer Cytoplasm manipulation, male sterility
Manipulation of medium composition Selection for growth, vigour, disease resistance, stress tolerance herbicide resistance, NPR response, in-vitro chemical
mutagenesis
Radiation in vitro, and in vivo Elimination of linkage drag in cybids/hybrids/wide crosses, induction of chromosomal exchanges
DNA probes Disease detection, sexing, trait selection, QTL maps
Recombinant DNA Transgenesis, gene location, insertion/deletion/mapping
1268 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
Fig. 4. The major area of plant cell and tissue culture and its application.
4.1. Tissue culture
Tissue culture comprises a set of in vitro techniques, methods
and strategies of biotechnology. This in vitro technique has been
applied for the diversification of targeted crops for improvement.
The improvement is in the form of the health of plant materials
and/or the availability of desired germplasms to plant breeders.
The area of plant tissue culture is vast (Fig. 4). Generally, callus
culture, cell suspension culture, protoplast culture, anther or
microspore culture and organ or meristem cultureare the areas
that can be observed in terms of practical aspects.
Less genetic diversity, poor seed germination, scanty and
delayed rooting of seedlings and vegetative cuttings demand the
practice of different forms of tissue culture to create more diverse
plants, inter-species hybridisation, and disease and virus free
plants for Jatropha improvement [9,41,83]. Moreover, the devel-
opment of an efficient regeneration system amenable to genetic
transformation is a prerequisite for plant genetic engineering.
Tissue-culture protocols are available for most crop species,
although continued optimisation is still required for many crops,
especially cereals and woody plants. For Jatropha tissue culture,
organogenesis and callus culture have been investigated by dif-
ferent authors (Table 4). Other areas, i.e., protoplast culture and
fusion, anther and microspore culture, ovary culture, and endo-
sperm culture, need to be investigated for variation creation and
breeding.
4.2. Organogenesis
De novo formation of organs (shoots or roots) by complex
process is called Organogenesis. Shoots can be derived either
through differentiation of non-meristematic tissues or through
pre-existing meristematic tissues. Appropriate choice of explant,
age of the explant, definite media formulations, specific growth
regulators, genotype, source of carbohydrate, gelling agent make
successful of organogenesis. Other physical factors including light
regime, temperature, humidity, etc are also important. Phyto
hormones are the regulator plants growth and development.
Appropriate ratio of auxins and cytokinins is important to induce
cell division and growth in plant tissue cultures.
Use of MS medium [84] is very common in vitro cultures, but
Warakagoda and Subasinghe [85] used B5 medium [86]. Different
parts of plant i.e. leaf [33,87–94], nodal [95], axillary nodes [88],
shoot tips [87,96,97], hypocotyl [87,98,99], epicotyl [98,100], and
cotyledons [101] are used as explants for tissue culture (Table 4).
The hormones TDZ, BAP, IBA, and NAA are used commonly.
There are some recent reports on micro-propagation from leaf
discs. Leaves from both in vitro and in vivo plants were used as
explants. Ethanol (70% V/V) and 0.1 % HgCl2 (W/V) were used for
explants sterilisation but the treatment duration varies depending
on explants types and sources. Culture condition was maintained
as temperature 25 72 °C, 16 h lighting of 25–57 mmol m  2s  1
white fluorescent cool light. Sing et al. studied the indirect orga-
nogenesis from leaf and petiole callus. It was observed that in vivo
leaf response less to hormones and media [102]. The same result
was observed by others [99,101,103]. Sing et al. reported MS
medium supplemented with BAP (4.44 μM) and IBA (2.45 μM) is
the best among studied media composition for regenerative callus
induction from petiole. Higher percentage of callus (91%) was
regenerated from petiole callus wearers 61% was from leaf callus
on media formulation with BAP (8.88 μM), IBA (0.49 μM), and ABA
 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277 1269

Table 4
Tissue culture studies of Jatropha.

Regeneration mode Explant Hormone(s) dose References

Callus mediated shooting Petiole 8.88 μM BAP, 0.49 μM IBA , and 1.9 μM ABA [102]
Shoot induction Node 1.5 mg/L BAPþ0.5 mg/L NAA [170]
Somatic Embryoenesis Leaf 0.5mg/L 2, 4-D and 5mg/L BAP [94]
Shoot induction Leaf 1.0 mg/L TDZ, 0.5 mg/L Kn, and 0.5 mg/L GA3. [93]
shoot induction Cotyledon 2.27 μM TDZ (shoot induction) 10 μM (Kn), 4.5 μM (BAP) and 5.5 μM ( shoot proliferation) [171]
shoot induction Hypocotyls 0.5 mg/L TDZ (shoot induction)2 mg /L Kn and 1 mg /L, BAP (elongation) [99]
shoot induction Petiole Non toxic 2.27 μM TDZ (shoot Induction) 10 μM Kn, 4.5 μM BA, and 5.5 μM NAA ( shoot proliferation) [103]
Shoot induction Cotyledon 3 mg/L BA and 0.1 mg/L IBA [92]
Shoot induction Leaf 0.90 μM TDZ and 0.98 μM IBA [90]
Shoot induction Petiole TDZ μM, 10 μM Kn, 4.5 μM BAP,5.5 μM NAA, [103]
Callus mediated shooting Immature embryo 1 mg/L BAþ0.5 mg/L Knþ 0.25 mg/L IBAþ500 mg/L PVPþ 30 mg/L citric acid [107]
Direct and indirecr organogenesis Epicotyls and hypocotyls 0.25 mg/L TDZ, 0.5 mg/L Kn, 0.25 mg/L IBA [98]
Direct organogenesis Cotyledonary leaf 9.08 μM TDZ, 10 μM Kn,4.5 μM BAP, 5.5 μM NAA [101]
Advantisious shoot bud Leaf disk 2.27 μM TDZ, 2.22 μM BAP, 0.49 μM IBA [89]
Direct organogenesis Node 22.2 μM BA, 2.3 μM Knþ 0.5 μM IBAþ (27.8–55.6 μM) Adenine sulphate [95]
Somatic Embryoenesis Leaf disk (2.3–9.3 μM) Kn, (0.5–4.9 μM) IBA, 13.6 μM Adenine sulphate [108]
Callus and Suspension culture Leaf and Hypocotyl 0.5 mg/L 2,4-D þ2% coconut milk [172]
Multiple soot induction Nodal segments 1.5 mg/L BAP, 0.5 mg/L Kn, 0.1 mg/L IAA [105]

(1.9 μM). Induced shoots were proliferated and elongated on BAP
(2.22 μM) and IAA (8.56 μM) media. Finally it were rooted on
media with IBA (2.45 μM), NAA (0.54 μM), and 0.02% activated
charcoal. They reported 98% survival rate of in vitro plants in green
house [102].
Zhang et al. studied the factors for direct shoot regeneration from
mature leaves. While studying hormonal (TDZ, Kn and GA3) effects,
the statistical analysis, L9(34) orthogonal test showed that thidiazuron
(TDZ) and GA3 played key role in bud induction stage. Shoot induction
is proportional to TDZ concentration. However, shoots from higher
concentration of TDZ did not perform well at elongation and pro-
liferation stages [93]. Other researchers observed similar effect of TDZ
[99,101,103]. Addition of abscisic acid (ABA) with BAP and IBA influ-
ences shoot regeneration from leaf and petiole callus whereas GA3
with BAP and IBA did not shown any improvement to the regenera-
tion efficiency [102]. Sharma et al. studied the Effect of Ca2 þ con-
centrations in the germination medium for shoot buds induction.
They found that the hypocotyl explants from the seedlings that was
raised in medium containing half strength Ca2 þ concentration
(0.22 g mL 1) and cultured on the TDZ supplemented medium
showed increased response. The Ca2 þ concentration is dispropor-
tionate to shoot induction. However, the absence of Ca2þ in the ger-
mination medium led to 100% mortality of explants [99]. Genotypes
and age of explants significantly affect direct organogenesis
[93,99,101]. Young leaves are more responsive to hormones also more
susceptible to surface sterilisation. Leaf discs placement (abaxial and
adaxial) on media is also a factor. Zhang et al. reported the best
medium among the studied hormonal combination for shoot bud
induction was Murashige and Skoog (MS) medium supplemented
with 1.0 mg/L TDZ, 0.5 mg/L Kn, and 0.5 mg/L GA3. The induced
shoots were proliferated and elongated on medium with 0.3 mg/L 6-
benzylaminopurine and 0.01 mg/L indole-3-butyric acid (IBA) and
finally rooted on half-MS media containing 2.0 mg/L IBA. It is also
reported that genotype affects regeneration frequency and the third to
fifth leaves from the apical bud were the best sources of explants [93].
Kumar et al. reported thidiazuron (TDZ) has greater influence on
regeneration as compare to BAP. They did subculture the induced soot
on MS media fortified with 10 μM kinetin (Kn), 4.5 μM BAP, and
5.5 μM α -naphthaleneacetic acid (NAA) and 2.25 μM BAP and 8.5 μM
IAA for shoot proliferation and elongation. The best rooting was from
hormone free half MS media with 0.25 mg/L activated charcoal after
four (4) days shock treatment on MS media with 15 μM IBA, 5.7 μM
IAA, and 11 μM NAA. It was found 90% successful plant establishment
on soil [101].
Plants regenerated from petiole retain genetic stability because,
it is driven from somatic cells of petiole [104]. Kumar et al. studied
organogenesis from petiole of non-toxic Jatropha and reported MS
media with 2.27 μM TDZ was best for shoot induction and future
proliferation. They also studied the orientation of placement of
petiole on media and found that shoot induction percentage and
number of shoot bud per explant were higher in case of horizontal
placement in compare to vertical placement. MS medium sup-
plemented with 2.25 μM BA and 8.5 μM IAA was used for shoot
elongation. And half-strength MS medium with 15 μM IBA, 11.4 μM
IAA and 5.5 μM NAA was used for in vitro rooting of elongated
shoots. Above 90% in vitro plants were successfully established on
soil [103].
Plants from in vitro node culture are genetically more stable
[104]. Kalimuthu et al. cultured nodal explants on MS medium
supplemented with 22.19 μM BAP, 2.32 μM Kn and 0.57 μM IAA for
shoot induction [105]. Shrivastava and Banerjee reported shoot
regeneration from axillary nodes using MS media in combination
4.90 μM IBA and 13.3 μM BAP along with growth additives (ade-
nine sulphate, glutamine and L-arginine) [106]. Whereas Datta
et al. observed that MS medium in combination with 22.19 μM BAP
and 108.58 μM adenine sulphate is suitable for axillary shoot bud
proliferation from nodal explants [95]. Rajore and Batra used MS
medium with BAP and IAA along with glutamine and adenine
sulphate for best response of shoot proliferation from shoot tip
[96]. Datta et al. used MS þ4.90 μM IBA for rooting of induced
shoot bud [95].
Some authors reported callus mediated regeneration from
immature cotyledons and embryos [107] and mature cotyledonary
leaf [87,90] of Jartopha. Varshney and Johnson observed the best
response when the explant (immature cotyledons and embryos)
were cultured on the medium containing IBA with BAP and CuSO4
[107]. Other growth additives did not have significant impact on
morphogenic callus formation.
The first complete protocol of somatic embryogenesis of
Jatropha was reported by Jha et al. [108]. They are able to produce
embryonic calli from leaf explants on MS medium with only
9.3 μM Kn. They also observed MS medium with 2.3 μM Kn and
1.0 μM IBA is the most effective combination for somatic embryo
development of Jatropha curcas and addition of 13.6 μM adenine
sulphate stimulated the process of development of somatic
embryos. Kalimuthu et al. was able to generate somatic embryos
on the surface of cotyledonary. They used MS medium containing
8.87 μM BAP. Somatic embryogenesis was also studied by
1270 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
Medipally et al.. They produced green embryonic callus on MS
media with BA (0.5% mg/L) and 2, 4-D (0.5 mg/L) [94].
4.3. Haploid plant production
Haploid plants and/or germaclonal variation can be produced
by anther, isolated microspore culture and ovary or ovule culture.
The term androgenesis refers to the culture of the anther, whereas
gynogenesis is the culture of the ovary or ovule. Embryos can be
developed directly without forming a callus or proliferation of the
callus from microspores of the anther. The underlying principle of
androgenesis is to stop the development of the pollen cell whose
fate is normally to become a gamete and to force its development
directly into a plant. Genotypes, the physical status of the donor
plant, the stage of pollen, pre-treatment of the anthers, and the
culture media composition are the factors of fruitful androgenesis.
Homozygous diploid plants can be produced by treatment (col-
chicines or endomitosis) of haploid plant. The potentials of haploid
plants in breeding are i) the development of pure homozygous
lines, ii) hybrid development, iii) induction of mutations, iv)
induction of genetic variability, v) generation of exclusive male
plants, vi) cytogenetic research, vii) significance in early release
variety, viii) hybrid sorting in haploid breeding, ix) disease resis-
tance, x) insect resistance, xi) salt tolerance and xii) double hap-
loids genome mapping [109] . The application of haploid breeding
for desired traits is proven. Some varieties of rice, wheat, and
tobacco are produced through anther culture (Table 5). High levels
of management and expertise, the production of triploid along
with diploid varieties, selective cell division, induced anabolism,
and doubling of haploid plants that do not allow production of
homozygotes are critical for haploid breeding.
4.4. Embryo culture
An embryo forms after successful fertilisation. Aseptic isolation
of either immature or mature embryos and in vitro culture for
obtaining viable plants is called embryo culture. Improvement of
the quality and yield potential of all major crops have been
achieved through interspecific and intervarietal crosses and
selection [110]. Wide hybridisation, where individuals from two
different species of the same genus of different genera are crossed,
often leads to failure. Pre and post-fertilisation barriers prevent
the successful transfer of genes from the wild to cultivars. Embryo
rescue can minimise the obstacles. There are reports of traits
transfer to cultivars through embryo rescue (Table 6). The selec-
tion and formulation of complex culture media is the most
important and critical aspect of immature embryo culture. Other
factors include isolation of the embryo without being damaged
and an embryo development stage during isolation [111]. Pre-
vention of embryo abortion in wide crosses and early ripening
stone fruits, haploid production, overcoming seed dormancy, and
Table 5
Crops varieties released through anther culture.
Crop Variety
Wheat Jinghua No. 1, Florin, Huapei 1, Lunghua 1, Delibab, GK Ambitus,
Jingdan 2288, Zing Hua 1, Zing Hua 3, Zing Hua 5
Tobacco Tan Yuh No. I, F 221, Tanyu-2, Tanyu-3, Lynd, Hai Hua-19, Hai Hua-29,
Hai Hua-30, Hai Hua-31
Rice Hua Uii 1, Hua Uii 2, Xin Xiu 1, Tan Feng No. 1, Ou-Hwa No. 1, Ou-Hwa
No. 2, Huayu-I, Huayu-2, Thong Hua 8,Thong Hua 9, Huajian 7902,
Huahanzhao, Nanhua 5, Nanhua 11, Oianhua 1, Yingyou No. 2, Hwa-
seongbyeo, Huayu 15, Zhe Keng 66, Hwajinbyeo Suweon 384, Hwa-
cheongbyeo, Shan Hua 369, Aya, Shirayukihime,Hwayeongbyeo, Hir-
ohikari, Hirohonami
Table 6
Resistant traits transfer to cultivar species through embryo rescue.
Crop species Resistance trait(s)
Lycopersicon esculentum * L. Virus, fungi, and nematodes
peruvianum
Solanum melogena * S. Brinjal shoot and fruit borer (L. arbonalis)
khasianum
Solanum tuberosum * S. Potato leaf roll virus
etuberosum
Brassica olerasia * Brassica napus Triazine resistance
B. napus * Brassica olerasia (kale) Cabbage aphid
Triticul aestivum * Thinopyrum Salt tolerent
scirpeum
Hordeum vulgare *Hordeum Powdery mildew
bulbosum
H. bulbosum * Hordeum vulgare Powdery mildew and spot blotch
Oryza sativa* Oryza minuta Blast (Pyricularia grisesa) and Bacterial blight
(Xanthomonas oryzae)
shortening of the breeding cycle are potential applications of
embryo culture in plant breeding. This study makes use of
embryology, in addition to the applied uses. Nutrient require-
ments, metabolism, growth requirements, phytohormones effects,
environmental conditions on zygotic embryogenesis and the
regeneration potentials of whole embryos and their segments can
be observed by culturing embryos outside the ovule. Promoters
and inhibitors of embryogenesis can be distinguished by this
technique [112].
4.5. Endosperm culture
Triploid endosperm is formed by the fusion of two polar nuclei
and one of the male gametes during the fertilisation process. Its
function is to support the developing embryo and its differentia-
tion. The endosperm is an excellent experimental system for
morphogenic studies because it lacks any differentiation, and it
consists mostly of parenchymatous cells. Triploid plants are in
commercial use for a number of crop species (e.g., banana, apple,
beet, tea, cassava, and mulberry) [113–118]. Conventionally, tri-
ploidsare produced by crossing between tetraploids and diploids.
The cross may not be successful due to problems associated with
the cross. Triploid plants can be raised through endosperm culture.
It is desirable to plant breeders to apply the techniques to eco-
nomically important plants where conventional breeding methods
prove futile. Both mature and immature endosperm culture are
reported for some species. Triploid rice, apple, citrus, santalum,
and croton are produced from immature endosperm culture.
Though in other species, such as Ricinus communis, Zea mays and
cucumis, unlimited callus growth is found, but organogenesis does
not occur [109]. The endosperm can also be used as nurse tissue
for raising hybrid embryos.
4.6. Somatic hybridisation
Species boundaries limit interspecies sexual hybridisation in
crop improvement programmes. Protoplast isolation, culture and
fusion from taxonomically distinct plants are the techniques for
the production of somatic hybrids. One of the major applications
of somatic hybridisation is the production of novel interspecies
and intergenic crosses. New genus can be produced by inter-genus
crossing (Table 7). Somatic hybridisation is also applicable for
potential gene transfer (Table 8). Many disease resistance genes,
including potato leaf roll virus, leaf blight, and Verticillium, phy-
tophthora resistant genes and abiotic resistant genes, have been
transferred to Solanum tuberosum from other species [119–121].
Blackleg disease resistance, resistance against nematodes and
 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277 1271

Table 7 Table 8
Intergenic protoplast fusion. Genetic traits transferred via protoplast culture.

Species 1 Species 2 New genous Refs. Crop Cultivar parent Donner parent Desired traits

Raphanus sativus Brassica oleracea Raphanobrassica [173] Tobacco Nicotiana tabacum N. nesophila Tobacco mosaic
(2n¼18,rr) (2n ¼18, cc) Virus
Moricandia arvensis Brassica oleracea Brassicomoricandia [174] Nicotiana tabacum N. repanda Tobacco mosaic
Eruca sativa (2n¼ 22) B. napus (2n¼ 38) Erucobrassica [175] Virus
Oryza sativa (2n¼ 24) (Echinochloa oryzicola Oryzicola [176] Nicotiana tabacum N. nesophila Tobacco horn worm
(2n ¼24) N. tabacum Nicotiana rustica High nicotine
Arabidopsis thaliana Brassica campestris Arabidobrassica [177] content
(2n¼10) (2n ¼20) N. tabacum N. sylvestris Streptomycin
Datura innoxia (2n¼ 48) Atropa belladonna Daturotropa [178] resistance
(2n ¼24) N. tabacum N. sylvestris CMS
Solanum tuberosum L. esculentum (2n ¼24) Solanopersicon [109] Brassica Brassica napus B. nigra Phoma lingam
(2n¼24) B. oleracea Sinapis alba Alternaria brassiceae
Nicotiana tabacum L. esculentum (2n ¼24) Nicotiopersicon [109] B. oleracea sp. B. oleracea (Ogura Cold tolerant
(2n¼24) capitata CMS line)
B. oleracea Brassica napus Black rot
B. nigra Brassica napus Hygromycin
Alternaria brasssiceae of B. napus have been developed [122]. resistance
B. campestris Brassica napus CMS
Tomato varieties resistance against TMV, potted wilt virus, insects,
Brassica napus B. tournefortii CMS
pests and cold tolerance are also reported. Improved quality Solanaceae S. tuberosum S. brevidens Potato leaf roll virus
characteristics, such as the low concentration of erucic acid con- S. tuberosum Solanum phureja Potato leaf roll virus
tent in Brassica and high content of nicotine in tobacco, are S. circalifolium S. tuberosum Phytophthora
S. melongena S. sanitwongsei Pseudomonas
reported. Cytoplasmic male sterility (CMS) lines of broccoli, sun-
solanacearum
flower, rice, and mustard of desired traits are produced by proto- S. tuberosum S. commersonii Frost tolerant
plast culture. Autotetraploids can also be produced. Hybridisation S. melongena S. sisymbrifollum Nematod
is possible at juvenile stages, so it can reduce generation intervals. S. melongena S. bulbocastanum Nematod, Root knot
Nematod
S. nigrum S. tuberosum Triazine resistance
4.7. Mutagenesis in crop improvement S. tuberosum S. chacoense Potato virus X
Tomato Lycopersicon L. peruvianum TMV, spotted wilt
Any heritable change in genetic material is called a mutation. esculentum virus, cold tolerant
Mutation can occur naturally or be induced by humans for specific Inter species S. lycopersicoides L. esculenttum Cold tlerent
S. ochranthum L. esculenttum Tomato disease
interests. Physical agents (e.g., UV, gamma, and X-ray), chemical insect pest
agents (e.g., EMS, NaN3, colchicines, herbicides, salinity, silver Citus sinensis Poncirus trifoliata Phytphthora
nitrate), and plant growth regulators (e.g., GA, IAA, BAP, JA) cause Raphanus sativa B. oleracea Var. Club rot disease
mutations in plants. Mutagenesis technology may produce botrytis
B. oleracea Var. S. alba þB. carinata Alternaria brassicola
favourable variants of wanted traits [123,124]. Mutagenesis can
botrytis and Phoma lingam
alter plants from germination to harvesting by the alteration of Citrullus lanatus Cucumis melo club rot disease
metabolism, tissues and organs. The interest in mutagenesis has Hordeum vulgare Daucus carota frost and salt
been increasing in plant breeding. These techniques have been resistance
Raphanus sativus Brassica napus Beet cyst nematode
executed widely for the improvement of crop yield, quality, dis-
Sinapis alba Raphanus sativus þ- Beet cyst nematode
ease, and pest resistance [125,126]. In vitro mutagenesis has been Brassica napus
used regularly to improve quality, resistance traits and innovative Brassica napus Eruca sativa low erucic acid
germplasm emerging [127]. A mutant varieties database reveals Brassica napus þB. R. sativa CMS and Triazine
campestris resistance
over 3200 released mutant varieties from 214 different plant
Brassica napus Sinapis alba Alternaria brassiceae
species in more than 60 countries throughout the world, among
which oilseed crops and oilseed rape comprise at least 63 and 25
varieties, respectively (http://www-infocris.iaea.org/MVD/). culture medium, and age of the culture are some factors affecting
The combined use of mutation with in vitro culture technology the types and frequency of somaclonal variation. The exact basis of
can be regarded as “an ideal system” for crop improvement. First, somaclonal variation is not clearly understood. It is assumed that
haploid or doubled haploid plants produced through culturing the possible causes of somaclonal variation include: (1) karyotypic
techniques can express all mutations, both recessive and dominant. changes, (2) point mutations, (3) somatic crossing over and sister
Therefore, recessive mutants can be detected in the first generation,
chromatid exchange, (4) somatic gene rearrangement, (5) DNA
and mutants can be fixed rapidly. Second, the probability of detec-
amplification, (6) transposable elements, (7) DNA methylation,
tion of beneficial mutants is high because it provides a large
(8) changes in organelle DNA, and (9) epigenetic variation. The
population available for mutation processes. Third, mutants and
major applications of somaclonal variation in crop breeding are
chimerism are avoidable during the production process.
Inereasing Genetie Variability for Agronomie Traits, Development
of 'Elite' Germplasm and Commercial Cultivars,and Enhancing
4.8. Somaclonal variation
Alien Gene Introgression into Cultivated Specie. A large number of
Genetic variability is crucial for plant breeding. Variation cre- plant species, e.g., potato, sugarcane, tobacco, tomato, wheat, rice,
ated by cell cultures, regenerated plants and their progenies is and brassica, for various agronomic traits, such as disease resis-
known as somaclonal variation [128]. Somaclonal variation can be tance, plant height, tiller number, maturity and for various phy-
created by callus growth, regeneration, in vitro screening, inher- siological and biochemical traits, have undergone Somac1onal
itability of the desired traits, and multiplication of stable variants variation. Some useful somac1onal variants have been obtained,
to develop new breeding lines. The explant source, genotype, and some of them have been released as cultivars (Table 9).
1272 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277

4.9. Genomic resources- DNA markers, marker assisted selection and programmes [129,130]. Prospective applications (Fig. 5) of these
molecular breeding genomic resources can be grouped according to Yue et al.
i) interim: during a 1–2 year period, including evaluation of
Genomic resources- DNA markers, linkage maps, genome genetic diversity, population and parent age relationship and bar
assembly data, express sequence tags (ESTs), and QTLs, can assist coding of elite trees, ii) middle term: during a 2–4 year period
in high quality and quantity Jatropha oil development linkage and QTL mapping and marker-assisted selection (MAS) and
producing transgenic trees for desired traits may be the applica-
Table 9 tion of genomic resources, iii) long duration: during this time (44
Target traits selected by somaclonal variation.
years) identification of a large number of SNPs from genomic data,
Crop Interested trait(s) Refs. genome wide association studies (GWAS) and genomic selection
(WS) for improved Jatropha are the application of genomic
Rice Lysine content [179] resources [23].
Resistance to blast (DAMA) [180]
The variable dispersed sequences in genome that can be dis-
Dwarf, Iod ging resistant 10% higher yield than [181]
the parent variety (Hatsuyme) tinguished by molecular and biochemical techniques are called
Tolerance to salt [182] DNA markers. Germplasm characterisation, linkage and QTL
Wheat Resistance to Helminthosporium [183] mapping, and molecular breeding are some direct applications of
Tolerance to heatldrought stress (KS89WGRC9)' [184]
molecular markers [131].
Resistance to barley yellow dwarf virus (BYDV) [185]
from Thinopyrum sp (TC5, TC6, TC9)" Different molecular markers, such as Random Amplified Poly-
Tolerance to frost [186,187] morphism DNA (RAPD), Enhanced Random Amplified Poly-
Tolerance to salt [188] morphism DNA (ERAPD), Simple Sequence Repeat (SSR), Single
Potato Resistance to Fusarium oxysporum [189]
nucleotide polymorphism (SNP),Sequence Characterised Amplified
Resistance to Phytophthora infestans [190, 191]
Sugarcane Resistance to eyespot, Fiji disease and [192, 193] Region (SCAR), Amplified Fragment Length Polymorphism (AFLP)
Resistance to eyespot [194] and Inter-Simple Sequence Repeat (ISSR),were developed since the
Sweet potato Darker and stable skin colour (Scarlet)b [195] decoding of the Jatropha genome and can be used to accelerate
Tobacco Herbicide resistance [196, 197]
breeding by Marker Assisted Selection (MAS) and genomic selec-
Resistance to potato virus Y (NC744)" [198]
Resistance to blue mould (NC-BMR42, 90)" [199] tion (GS) for desired traits at a very early stage [132–134].
Maize Resistance to Helminthosporium maydis [200] The essential framework, linkage map, is the association
Tomato High solid (DNAP9)" Evans, [201] between DNA markers and traits. First generation linkage maps
Resistance to race 2 of Fusarium (DNAPI7) [201]
were constructed by Liu et al. SSRs and SNPs are the co-dominant
Celery Resistance to Fusarium wilt (UC-T3)" [202]
Brassica Herbicide resistance [203] markers that they used for genotyping. They used 96 individuals
Tolerance to salt [204] from two mapping population families. Interspecies (J. curcas  J.
Lathyrus sativus Low neurotoxin [205] integerrima) cross and backcross was performed for the develop-
Birds-foot Resistance to sulphonyl herbicide(H401-4-4-2)" [206]
ment of the families [135].
trefoil
Bell pepper Fruits with fewer seeds (Bell sweet)' [201] Poor diversity of Jatropha in nature is assumed in the region for
Bermuda grass Fall army worm resistance (Brazos-R3)" [207] the advancement of linkage mapping. This makes the generation
Red cIover Regeneration ability (NEWRC)" [208] of informative reference families for mapping difficult [135].
Sorghum Resistance to Fall army worm (GATCCP 100/101)" [209]
Therefore, inter-species hybridisation is the best option. Inter-
Tolerance to acid soil (GACI02)" (GCI03/104)" [210,211]
species hybridisation of J. curcas with Jatropha. integerrima,

Fig. 5. Application areas of genomic resources in improving J. curcas [23].

 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
Jatropha. canascens, and Jatropha. gossypifolia produces viable off-
spring [33].
Transcriptome is the term that includes all RNA molecules
(mRNA, tRNA, rRNA, miRNA and other non-coding long or short
RNA) in one or a population of cells. In different cells and tissue in
different environmental conditions, the transcriptome can vary.
For a better understanding of the function and biological process
of a genome, transcriptome study is crucial. Costa et al. recognised
the majority of known genes in lipid synthesis and the degrada-
tion pathway by sequencing 13,249 ESTs from developing and
germinating seeds [136]. Another finding was the recognition of
some ESTs that might contribute to seed toxicity. Chen et al. found
9844 unique sequences (1070 contigs and 3595 singletons) by
sequencing ESTs. Three cDNA libraries with mRNA from embryos
at different developmental stages were constructed for this work
[137]. Another full-length enriched cDNA library from developing
seeds was developed by Yadav et al. [138]. King et al. sequenced
(454 high-throughput sequencing) the transcriptome of develop-
ing seeds and generated 46 Mb of raw sequence data, including
95,692 sequences using a single sequencing run [139]. Purush-
othaman and Madasamy developed cDNAs from developing seeds,
flowers, roots, embryos and mature leaves of J. curcas. They per-
formed 454 pyro sequencings and generated 383,918 raw reads,
among which 381,957 reads were high-quality [140]. Silva et al.
generated high-quality sequences of 11.8 giga bases by sequencing
ESTs from polled RNA. They used two lanes of the Illumina
sequencing platform [141]. Some genes related to the fatty acid
biosynthesis pathway were recognised [142]. Gu et al. sequenced
over 50,000 EST clones by constructing several cDNA libraries from
different tissues using Sanger sequencing.
A quality inclusive reference transcriptome, including all tran-
scripts, coding and non-coding, large and small RNA, is necessary for
transcriptomics studies. However, researchers are still looking for a
complete reference transcriptome of J. curcas [143]. Therefore,
assembling all of the available EST data to obtain a quality assembly
and annotating the inclusive transcriptome of Jatropha is urgent.
4.10. Genetic engineering
The rearrangement of the genetic materials of plants and ani-
mals by molecular and cellular biological techniques is genetic
engineering. It has proven to have many uses for desired changes
in many crops. The enzymes diacylglycerol acyl transferase, lyso-
phosphatidic acid acyltransferase, and glycerol-3-phosphate acyl-
transferase are involved in the fatty acids biosynthesis pathway.
Over expression of these enzymes increased the seed oil content in
Brassica and Arabidopsis [144–146]. Mutations in the genes of 3-
keto-acyl-ACP synthase II increases the 16-carbon fatty acids and
decreases 18-carbon fatty acids. The use of the mutant 3-keto-
acyl-ACP synthase II gene changes the seed oil composition [147].
Stearoyl-ACP desaturase gene silencing has a significant effect on
the stearic acid (saturated fatty acid) content in Brassica [148].
Therefore, it is possible to increase the oil content and specialise
the seed oil composition in Jatropha.
The carboxyl transferase of the ACCase b subunit, the biotin car-
boxyl carrier protein of ACCase, malonyl-CoA:ACP transacylase, 3-
keto acyl ACP reductase, beta-keto acyl ACP synthase I, betaketo acyl
ACP synthase II and acyl carrier protein are involved in fatty acid
biosynthesis in plastids; ω-3-fatty acid desaturase and ω-6-fatty acid
desaturase are involved in desaturation of fatty acids; acyl ACP
thioesterase A is involvedin the hydrolysis of fatty acids from acyl-
ACP; long chain acyl-CoA synthetase and acyl -CoA binding protein
are involved in activation and transport of free fatty acids to the
endoplasmic reticulum; glycerol-3 phosphate acyl transferase and
lysophosphatidic acid acyl transferase are involved in serial incor-
poration of activated fatty acids to the glycerol backbone to form
1273
triacylglycerol or oil [149]. These genes of the beta oxidation pathway
can be a good source of clones for oil yield and property
manipulation.
Gene manipulation can create resistance plants. Abiotic stress
(salt) tolerant carrot, maize and tomato are reported by increasing
the expression level of the betaine aldehyde dehydrogenase gene
[150–152]. It is also reported that drought, heat and salt stresses
increase the glycine betaine level in Jatropha [153]. Pennisetum
glaucum showed increased levels of salt tolerance after increasing
the expression level of the Na þ /H þ antiporter in transgenic Brassica
[154], and Escherichia coli trehalose-6-phosphate synthasegene
overexpression showed drought and salt tolerance in rice [155].
Some stress related unigenes, including trehalose-6-phosphate
synthase, spermidine synthase, late embryogenesis abundant pro-
teins, glutathione peroxidase, glutathione s-transferase, phosphor
ethanolamine N-methyltransferase, Na þ /H þ antiporter, ethylene-
responsive transcription factors,aquaporin, and the salt tolerance
protein, ascorbate peroxidase are identified [149]. These genes are
potential sources of stress tolerance in transgenic plants.
Genetic engineering has proven contributions for crop
improvement. Many commercial varieties of many cultivars have
been released. Some are very good and are widely cultivated
throughout the world. For Jatropha, genetic engineering work is
just being started. Several articles [156–162] on the transgenesis of
Jatropha have been published very recently. More investigation is
needed on the transgenesis for more female flowers, lower toxicity
components, disease and pest resistance. Jatropha is non-edible, so
it is beyond the controversy of genetic modification (GM) as
concern of human health.
5. Conclusions
The world is looking for renewable energy sources to minimise
the dependence on fossil fuels and environmental pollution.
Among the oil seed crops as sources of biofuel, Jatropha is one of
the best candidates. Moreover, it does not compete with food
crops. However, it did not meet the expected performance due to a
lack of improved varieties, large scale propagation without eval-
uating the planting material, knowledge gap, and consideration as
low impute crop. A number of improvement programmes have
been launched, but the desired commercial variety is still
unavailable.
Germplasm collection, characterisation, and evaluation are the
basis of a breeding programme. To accumulate and use the spe-
cialised but scattered knowledge, an international organisation
can be formed by the collaboration of major Jatropha cultivators,
such as India, China, Malaysia, Indonesia, Brazil, Mexico, and South
Africa. The R&D expertise and the lessons learned by these coun-
tries during last several years of Jatropha cultivation can be shared
and used to design genetic and molecular breeding strategies.
Biological techniques, in vitro micropropagation, anther and
microspore culture, ovary and ovule culture, protoplast culture,
nucleolus culture, endosperm culture, and somatic embryogenesis
can be used for the production of somaclonal and germaclonal
variation. In vitro mutagenesis is another means of variation
creation. In vitro selection for desired traits is promising for crop
breeding. Some reports in the last few years have utilised only
micropropagation. Other in vitro techniques need to be investi-
gated because of their proven potential for crop improvement.
Some molecular work has been performed, and some devel-
oped genomic resources of Jatropha have been used for the
assessment of the genetic variation in the natural population,
linkage and QTL mapping. However, Jatropha genome is in
advanced in comparison to the model and other agricultural sys-
tems. A high density linkage map to determine the association of
1274 M. Moniruzzaman et al. / Renewable and Sustainable Energy Reviews 54 (2016) 1262–1277
markers with high oil yield is needed for researchers. The J. curcas
genome is very small (ca.400 Mb). Thus, marker assisted selection
(MAS), genome wide association studies (GWAS), and genomic
selection (GS) could be even more attractive. A well-assembled
references genome of Jatropha is indispensable for these applica-
tions. The advancement of sequencing technology has sharply
reduced the cost of genotyping by sequencing and has stimulated
GWAS and GS studies.
Stable integration of desired gene(s) has been applied for crop
improvement. Recently, a protocol for agro bacterium mediated
gene transfer to Jatropha has been published. Genes involved in
fatty acid biosynthesis have been identified. Priority should be
given to increasing female flowers in inflorescence, reducing tox-
ins and increasing resistance.
Acknowledgement
The authors thanked The National University, Malaysia (UKM)
for providing research facilities and fund. One of the authors also
thanked the government of the people republic of Bangladesh for
allowing study leave to work on it. This study would not have been
possible without the cooperation of the staff and practical students
of Livinglives and future crop centre, UKM. This project is financed
under grant DIP-2014-011.
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