J. For. Res.

(2019) 30(3):745–754
https://doi.org/10.1007/s11676-018-0714-6

REVIEW ARTICLE

Red sandalwood (Pterocarpus santalinus L. f.): biology,

importance, propagation and micropropagation
Jaime A. Teixeira da Silva1 • Mafatlal M. Kher2 • Deepak Soner3 •

M. Nataraj2

Received: 3 November 2017 / Accepted: 6 February 2018 / Published online: 20 June 2018
 The Author(s) 2018

Abstract Pterocarpus santalinus L. f. (Fabaceae; red Keywords Conservation
Fabaceae
IUCN red data list

sanders) is prized for its wood whose colour and fragrance Medicinal plant
Micropropagation
Red sanders
is due to the presence of santalins that have pharmaceutical
and industrial uses. Red sanders is listed as an endangered
plant species on the IUCN red data list as a result of the Historical, cultural, medicinal, and economic
exploitation of its wood and essential oil. This review importance of red sanders as a basis
emphasizes the pollination biology, seed germination, for conservation
vegetative propagation and micropropagation of P. san-
talinus. Excessive use of P. santalinus has also caused the Pterocarpus santalinus L. f. (Fabaceae) is most commonly
emergence of various adulterants, so accurate identification known as red sandalwood in English, but it also has other
is essential. common names in several languages (Table S1). Ptero-
carpus is derived from the Greek words pteron (wing) and
karpos (fruit), referring to the winged pod, while santalinus
originates from the Latin sandal and inus (meaning similar
The online version is available at http://www.springerlink.com
to), i.e., a plant with characteristics similar to Indian san-
Corresponding editor: Yu Lei. dalwood, Santalum album L. (Botanical Survey of India
2012). Like African or Nepalese sandalwood (Teixeira da
Electronic supplementary material The online version of this Silva et al. 2016a) and Indian sandalwood (Teixeira da Silva
article (https://doi.org/10.1007/s11676-018-0714-6) contains supple-
mentary material, which is available to authorized users. et al. 2016b), P. santalinus is also prized for its hard, dark-
purple, bitter heartwood (Navada and Vittal 2014). In India,
& Jaime A. Teixeira da Silva the natural range of P. santalinus used to be a very restricted
[email protected] area of 15,540 km2 in the southeast (Sarma 1993). Cur-
& Mafatlal M. Kher rently, P. santalinus is found exclusively in a well-defined
[email protected] forest tract of Andhra Pradesh in Southern India (Raju and
& Deepak Soner Nagaraju 1999; Prakash et al. 2006; Balaraju et al. 2011),
[email protected] but is also found in the Chinese provinces of Yunnan,
& M. Nataraj Guangdong and Guangxi, and on Hainan Island, where it is
[email protected] referred to as zitan (Kaner et al. 2013).
1
P. O. Box 7, Miki-cho post office, Ikenobe 3011-2,
The colour and fragrance of P. santalinus heartwood are
Kagawa-ken 761-0799, Japan derived from santalins while the pleasent aroma is caused by
2 the presence of terpenoids (Kumar et al. 1974). A dye pre-
P.G. Department of Biosciences, Sardar Patel University,
Sardar Patel Maidan, Vadtal Rd., P.O. Box 39, pared from the heartwood of P. santalinus is used as a stain in
Vallabh Vidyanagar, Gujarat 388120, India light microscopy (Banerjee and Mukherjee 1981; Sen Gupta
3
Resonance, Plot No - 21/5, Near Regional Science Centre, and Mukherjee 1981), as a coloring agent in pharmaceutical
Acharya Vihar, Bhubaneswar, Odisha 751013, India preparations, in food, leather and textile industries (Ankalaiah

123
746 J. A. Teixeira da Silva et al.

et al. 2017), and as a textile dye (Gulrajani et al. 2002). The Flowering is discontinuous, blooming at intervals of 2–5 d
medicinal properties of P. santalinus have been extensively (Rao et al. 2001). Flowers open at night and the primary
reviewed elsewhere (Navada and Vittal 2014; Azamthulla pollinators are Apis dorsata, A. cerana indica and A. florea
et al. 2015) and will not be covered in this review. However, (Rao and Raju 2002). P. santalinus, which shows facultative
multiple uses (Table S2), ethnomedicinal uses (Table S3), and xenogamy, tends to eliminate growing fruits from self-polli-
phytochemistry (Table S4) have been provided as supple- nated flowers, i.e., there is large-scale abortion of flower buds,
mentary tables to offer a more rounded appreciation of this flowers and fruit (Rao et al. 2001), and has very low fruit set
tree in the context of this review. (- 6%), 52% of which set seed (Rao and Raju 2002).
The texture and colour differentiate good quality from
poor quality trees, with ‘‘wavy grain wood texture with Seed germination
intense red color’’ in the former and ‘‘straight grain wood
texture with light red color’’ in the latter (Prakash et al. Traditional seed propagation of P. santalinus yields low
2006), and it is the superior quality of P. santalinus that germination percentages due to a hard testa, poor viability,
makes it popular in the furniture industry (Prakash et al. and sensitivity to temperature (Kumar and Gopal 1975;
2006; Arunkumar et al. 2011; Arunkumar and Joshi 2014; Dayanand and Lohidas 1988; Anuradha and Pullaiah 1998;
Azamthulla et al. 2015). In Japan, P. santalinus is used to Naidu 2001a, b; Naidu and Rajendrudu 2001). Dried,
make carvings and musical instruments, shamisen and koto soaked and scarified P. santalinus pods resulted in 49%
(Kukrety et al. 2013b; Arunkumar and Joshi 2014; Azam- germination (Kumarasinghe et al. 2003) although seed
thulla et al. 2015; Ramabrahmam and Sujatha 2016), as well germination in natural stands or under artificial propagation
as name seals or hankos. In Buddhism, P. santalinus is is generally low (- 30%) (Kumar and Gopal 1975; Day-
considered to be a symbol of holiness, and is thus used for anand and Lohidas 1988; Kalimuthu and Lakshmanan
carved statues, as a constituent of incense (Wu et al. 2011), 1995; Naidu 2001a, b;Naidu and Rajendrudu 2001).
and for cremation (Ramakrishna 1962). In China, P. san- Alternate wetting and drying every 48 h enhanced germi-
talinus wood has a long history of use in furniture and other nation, reaching 73% (Vijayalakshmi and Renganayaki
valuable wood products (Berliner 1996; Kaner et al. 2013). 2017). Seed germination, seedling height, and root collar
The export of P. santalinus from India to Europe started diameter were all significantly stimulated by fire (Kukrety
in the 17th century, mainly for fabric dyeing (Vedavathy et al. 2013b). Presoaking P. santalinus pods with 500 mg/L
2004). The Herbal Folklore Research Centre in Tirupati, gibberellic acid for 24 h resulted in 66.7% seed germina-
India, estimated that from 500 planted trees ha-1, at least tion, as well as improved plant growth and seedling sur-
500 kg of heartwood per tree can be obtained after vival relative to other treatments with tap water, luke warm
25 years, thus 25 t ha-1 of wood plantation (Vedavathy water, gibberellic acid, H2SO4 or HCl (Patel et al. 2018).
2004). At 2004 prices of Rs. 75 kg-1, such a plantation
would yield a return of Rs. 177.5 lakhs ha-1 Vegetative propagation
(US$375,000 ha-1) (Vedavathy 2004). Current market
prices are, however, unknown to the authors, although Vegetative propagation of P. santalinus by semi-hardwood
prices are likely to be high since natural P. santalinus cuttings, cleft grafting, or air layering is not able to produce
stands have been in decline as a result of this overex- stock numbers required for effective preservation or for
ploitation for commercial purposes, earning it an endan- commercial purposes (Kedharnath et al. 1976; Kesava
gered status since 1997 (IUCN 2018). Reddy and Srivasuki 1990). Relative to seed germination,
This review provides an overview of the reproductive there are almost no studies on ex vitro vegetative propaga-
biology, seed germination and micropropagation of P. tion for red sanders. However, to provide elite germplasm
santalinus as tools for its conservation and large-scale with desired traits, such as the wavy grain or phytochemicals
propagation. such as santalins, vegetative propagation under controlled
conditions is desirable, and in vitro propagation allows for
the production of true-to-type plants via micropropagation
Basic flowering biology, and sexual and vegetative such as axillary shoot multiplication or shoot tip culture at a
reproduction large scale, to continuously produce plantlets with uniform
characteristics. In tree biotechnology, such as for Indian
Pollination and seedset sandalwood (Teixeira da Silva et al. 2016b), in vitro prop-
agation also allows for the improvement of desired charac-
The P. santalinus tree flowers in the dry season (Rao et al. teristics such as pathogen resistance or improved wood
2001; Rao and Raju 2002). The flowers are papilionaceous, quality by genetic engineering. The next section assesses the
bisexual, large and and yellow (Rao and Raju 2002). progress of micropropagation of P. santalinus.

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Red sandalwood (Pterocarpus santalinus L. f.): biology, importance, propagation and… 747

Micropropagation santalinus (Anuradha and Pullaiah 1999a; Arockiasamy

et al. 2000; Rajeswari and Paliwal 2008; Warakagoda and
Explants Subasinghe 2013). BA, alone or combined with other
cytokinins or auxins, has frequently been utilized for the
The explant source (i.e., mother plant) and the procedure to micropropagation of P. santalinus (Table 2). Prakash et al.
surface disinfect explants are important aspects underlying (2006) cultured nodes of mature trees directly on filter
the success of a tissue culture protocol (Leifert et al. 1994; paper bridges employing liquid MS medium containing
Teixeira da Silva et al. 2016c). Information about the 1.16–9.30 lM kinetin or 1.11–8.88 lM BA, as well as an
explants used for the in vitro propagation of P. santalinus, antioxidant, observing that 4.44 lM BA was optimum for
as well as surface disinfection protocols, are summarised in bud break and shoot multiplication.
Table 1. P. santalinus seedlings derived from in vitro seed
germination have been a popular source of explants, while In vitro propagation (callogenesis, regeneration
in vitro germinated seedlings, shoot tips, cotyledons, and somatic embryogenesis)
hypocotyls, mesocotyl and nodes have also served as
popular sources of explants for culture initiation since they Leaf, cotyledon, root, internode and nodal segments (pre-
do not require surface disinfection (Lakshmi Sita et al. sumably with axillary buds) from in vitro P. santalinus
1992; Anuradha and Pullaiah 1999a,b; Chaturani et al. seedlings formed callus, but shoot regeneration was not
2005; Rajeswari and Paliwal 2008; Balaraju et al. 2011; reported (Chaturani et al. 2005). Callus was induced from
Vipranarayana et al. 2012; Warakagoda and Subasinghe leaves and internodes of P. santalinus by Ashrafee et al.
2013). In terms of ex vitro sources of explants, Prakash (2014) solely to assess antibiotic activity against Aero-
et al. (2006) used young terminal shoot cuttings collected monas and Pseudomonas but regeneration was not asses-
from mature trees in winter as the explant; Ashrafee et al. sed. Details of effective plant growth regulator
(2014) used leaf segments from 1 to 2 year old plants while concentrations and combinations, medium composition,
Sarita et al. (1988) used nodes and terminal cuttings. and explant type, as well as their effects on morphogenesis
are presented in Table 2.
Basal medium
Rooting and acclimatization
The most commonly used and preferred basal medium for
in vitro studies on P. santalinus is Murashige and Skoog Sucessful roooting of in vitro-raised plants followed by
(1962) (MS) medium (Table 2). Lakshmi Sita et al. (1992) effective acclimatization and successful transfer of in vitro-
used Gamborg’s B5 medium (Gamborg et al. 1968) with propagated plants to field conditions is the final objective
2% sucrose and 0.8% agar to multiply axillary shoots from of any micropopagation protocol and care is needed to
shoot tips derived from seedlings germinated in vitro. avoid hyperhydricity in in vitro-raised plants, which tend to
Chaturani et al. (2006) used Anderson medium (Anderson display poor rooting efficiency (Ruffoni and Savona 2013;
1980) and Vitis medium (Chee and Pool 1987) to germinate Teixeira da Silva et al. 2017b). Rooting and acclimatization
seed. Anuradha and Pullaiah (1999a) employed half- protocols for in vitro-raised shoots of P. santalinus are
strength B5 medium supplemented with 0.05% activated summarized in Table 2. Only a few studies have quantified
charcoal to germinate seeds in vitro then used B5 medium the survival of micropropagated plants (Lakshmi Sita et al.
with 8.88 lM 6-benzyladenine (BA) for shoot tip culture. 1992; Prakash et al. 2006; Rajeswari and Paliwal 2008;
Balaraju et al. 2011; Warakagoda and Subasinghe 2013).
In vitro propagation from predetermined meristems Among the 12 reports on P. santalinus tissue culture,
in vitro rooting employed full-strength, half-strength and
Three predetermined meristems were employed in P. quarter-strength MS medium (Table 2). According to
santalinus tissue culture: shoot tips, cotyledonary nodes Arockiasamy et al. (2000), quarter-strength MS medium
and nodes from mature trees (Table 2). Shoot tips from supplemented with 5.71 lM IAA was effective for 76.2%
in vitro germinated seedlings were used by Lakshmi Sita rooting of shoots derived from cotyledenory nodes, but
et al. (1992), Anuradha and Pullaiah (1999a), and Balaraju acclimatization and survival of plantlets were not reported.
et al. (2011) for shoot tip culture, with either BA as the Vipranarayana et al. (2012) applied a pulse treatment of
most effective cytokinin (Lakshmi Sita et al. 1992; Anu- 7.34 lM IBA in half-strength MS but details about survival
radha and Pullaiah 1999a) or a combination of BA and were not reported. Rajeswari and Paliwal (2008) achieved
thidiazuron (Balaraju et al. 2011). Cotyldenory nodes were 85% plantlet survival ex vitro after a pulse treatment of
sucessfully applied for the in vitro propagation of P. 5 lM IAA and 1 lM IBA for 25 d. In contrast, Warak-
agoda and Subasinghe (2013) showed limited success (46%

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748 J. A. Teixeira da Silva et al.

Table 1 Explant source, size and surface sterilization procedures for preparation of tissue culture studies of Pterocarpus santalinus (chrono-
logical listing)
Explant source Explant type, size and density; culture Surface sterilization and preparation References
vessel

Seeds (soaked and dried for 15 d) ? Size of shoot tips (15-d-old seedling) Shoot tips: RTW (duration NR) ? soap Lakshmi Sita
seedlings. Age and source of mother NR. 5–8 mm shoot tips and nodes (soap name, duration of treatment and et al. (1992)
plant NR from in vitro grown shoots. Test tubes concentration NR) ? 0.1% HgCl2
(15 mL/tube) 15 min ? 3-4X SDW
Seeds from the wild ? seedlings. Age Roots, hypocotyls, mesocotyls, Pods: 50% HCl ? 50% EtOH 2–3 h ? Anuradha and
of mother plant NR cotyledons, shoot tips, nodes, leaves, TRW ? dried for 2 d ? pods opened Pullaiah
internodes (size NR for all explants) and seeds sown directly in vitro (1999a)
from 15-d old seedlings 6–7 cm tall
with 3–4 nodes. Test tubes (1
explant/tube)
Seeds from the wild ? seedlings. Age Hypocotyls (1 cm), epicotyls (1 cm), Seeds from dried pods: 70% alcohol Arockiasamy
of mother plant NR cotyledons (1.5 cm), shoot tips 1 min ? 0.1% HgCl2 ? 0.1% sodium et al. (2000)
(0.5 cm), internodes (0.5 cm), axillary dodecyl sulfate 10 min ? 5X DDW
nodes (0.5 cm) from 20-d old ? SDW 24 h ? SDW replaced every
seedlings. Test tubes (1 explant/tube) 8h
Pods (\3 to [5 cm) including wing. Seeds from pods of various sizes. Pods Seeds: 5% Teepol (duration NR) ? Chaturani et al.
Age of mother plant NR stored at 28 ± 5 C for 1–4 weeks. RTW 2 h ? 0.1% HgCl2 ? 2 drops (2006)
Culture vessel NR Tween-20 (5–25 min) ? 3X SDW
10 min
10-y-old tree, sampled in Nov.-Jan. Mature nodes from terminal shoots Shoots: 1% Teepol 30 min ? cut into Prakash et al.
(7–8 cm long). 25 9 150 mL test nodal segments 2–3 cm long ? 70% (2006)
tubes (1 explant/tube) EtOH 2 min ? 0.1% HgCl2 ? 0.1%
Tween-20 7 min ? 5X SDW
Mature pods and nodes (forest and Pods scarified in boiling water (5 min) Seeds and nodes: RTW 30 min ? testa Padmalatha
campus culture) or 5% H2SO4 (10 min). Culture vessel removed ? 2% Bavistin (15 min for and Prasad
NR nodes, 30 min for seeds) ? 70% (2007, 2008)
EtOH 2 min ? 0.1% HgCl2 (15 min
for nodes, 12 min for seeds) ? 4-5X
SDW
Seeds ? seedlings. Age and source of Seedling-derived cotyledonary nodes Explants from 30-d-old seedlings: RTW Rajeswari and
mother plant NR (1.5 cm long), nodal segments (1 cm 1 h ? 0.025% Tween-20 10 min ? Paliwal
long). Test tubes (1 explant/tube) 3X DW ? 0.1% HgCl2 10 min ? 3X (2008)
SDW
Seeds from the wild ? seedlings Shoot tips of 20-d-old in vivo seedlings. Peeled seeds: 1% Bavistin (fungicide) Balaraju et al.
50-mL Borosil test tubes (1–2 10 min ? wash with H2O ? (2011)
explants/tube) 50–300 ppm GA3 24 h. Apical
meristem explants: RTW 10 min ?
DW ? some drops Tween-20 5 min
? SDW 2-3X ? 1% Bavistin 5 min
? SDW ? 70% EtOH 30 s ? SDW
2-3X ? 0.1% HgCl2 3 min ? 4X
SDW
In vitro seedlings (age NR) Nodal segments; one explant per test Seeds: RTW 30–40 min ? 5% Teepol- Vipranarayana
tube (150 mm 9 25 mm) B-300 (wetting agent) stirring 15 min et al. (2012)
? 1% Bavistin Carbandazim
(fungicide) 10 min ? DW 30 min ?
70% EtOH 30 s ? 0.05% HgCl2
5–10 min ? 4X SDW

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Table 1 continued
Explant source Explant type, size and density; culture Surface sterilization and preparation References
vessel

1. Seeds from green-brown pods from 1. Mesocotyl segments, cotyledonary Seeds: 10% Clorox (fungicide) 20 min Warakagoda
25-y-old trees nodal segments, shoot tips, from 20-d- ? 70% EtOH 2 min ? rinse NR and
2. Age of mother plants in greenhouse old in vitro germinated seedlings and Shoot segments: 5% Teepol 5 min ? Subasinghe
NR. Greenhouse conditions NR. seedlings RTW 30 min ? 0.3% topsin solution (2013)
Terminal bud removed and sprayed 2. Immature and semi-hard shoot 1 h ? 2X SDW ? Clorox ? 2 drops
with 10 mg/l BA at 2-w intervals. 70% segments Tween-20 (10, 15, 20% for 10, 15,
thiophanate methyl (topsin) sprayed at Culture vessel and other conditions for 1 20 min each) ? 70% EtOH 2 min ?
100 mg/l 24 h before collecting and 2 NR 2X DW. Whole process repeated twice
explants. Plants treated with 200 mg/l
Albert’s solution (liquid fertilizer) at
2-w intervals
1–3-y-old trees Leaves and internodes. Size and density Leaves: wash in DW ? 70% EtOH 30 s Ashrafee et al.
NR; test tubes ? 0.1% HgCl2 3 min. Internodes: (2014)
70% EtOH 2 min ? 0.1% HgCl2 ?
2–3 drops Tween-20 7 min. No rinses
described for both explants
d day(s), DW distilled water, DDW double distilled water, EtOH ethyl alcohol (ethanol), GA3 gibberellic acid, HgCl2 mercury chloride, IZE
immature zygotic embryo, NaOCl sodium hypochlorite, NR not reported in the study, RTW running tap water, s second(s), SDW sterilized (by
autoclaving) distilled water, SW sterilized water, y year(s)

of plantlets rooted) ex vitro with 49,000 lM IBA, possibly Woody anatomy such as grain waviness can be used to
because of the excessively high concentration of this auxin. delimit and identify P. santalinus (Rawat and Uniyal 1996;
Gasson and MacLachlan 2010). The Botanical Survey of
India (2012) used various anatomical methods such as
Variability in quality and quality control maceration, scanning electron microscopy, exo- and
endomorphic features, and fluorescence analysis to cor-
There is a problem with the adulteration and falsification of rectly identify P. santalinus wood samples.
plant material in the P. santalinus market. The heartwood Molecular markers are regularly utilized to measure the
of Adenanthera pavonina Willd. (Mimosaceae), known as degree of genetic variation within natural or breeding
‘Ranjana’ and ‘Raktakambal’ in West Bengal and ‘Bari populations, and have been extensively used in Indian
Gumchi’ in the northern parts of India, is often sold as a sandalwood research (Teixeira da Silva et al. 2017a). In P.
fake substitute for P. santalinus, while artificially colored santalinus, RAPD (random amplified polymorphic DNA)-
wood shavings and the sawdust of some other trees are also based marker analysis was used to detect variations in
sold on the market as cheap substitutes (Botanical Survey micropropagated plants raised from shoot tips, verifying
of India 2012). In China, the manufacture of furniture that in fact no variation existed (Balaraju et al. 2011).
utilizes Dalbergia louvelii R. Vig. (violet rosewood) as a RAPD was also used by Usha et al. (2013) to detect vari-
substitute for P. santalinus since both plants have a very ation among nursery-grown plants. Variation in genetic
similar appearance and anatomical characteristics, and distance among natural accessions, detected by RAPD
cheaper D. louvelii is often illegally used to impersonate markers, reflected a high level of DNA polymorphism due
the valuable P. santalinus (Zhang et al. 2014). Zhang et al. to outcrossing (Padmalatha and Prasad 2007; Usha et al.
(2014) used conventional infrared spectroscopy (FT-IR), 2013). Jhansi Rani and Usha (2013) developed a sequence
second derivative infrared (SD-IR) spectroscopy and two- characterized amplified region (SCAR) marker to differ-
dimensional correlation infrared (2DCOS-IR) spectroscopy entiate wavy from straight-grained plants at the seedling
to differentiate furniture made of P. santalinus wood from stage. Jyothi et al. (2014) reported differences in the
furniture made from D. louvelii. They observed that P. quantity of genomic DNA in samples collected from dif-
santalinus wood had a higher holocellulose content than D. ferent locations in Andhra Pradesh, India.
louvelii wood while D. louvelii had more NaOH- and Therefore, quality control, as assessed by anatomical or
benzyl-alcohol-based extracts than P. santalinus. chemical methods, is essential to verify the originality of P.
The size and age of trees affects the heartwood content santalinus wood while molecular methods serve to confirm
and wood density of P. santalinus (Suresh et al. 2017). genetic stability.

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Table 2 In vitro conditions for tissue culture studies of Pterocarpus santalinus (chronological listing)
Culture medium, PGRs, Culture conditionsa Experimental outcome, maximum productivity, acclimatization References
additives, subcultures and variation

B5 ? 0.88 lM or 4.44 lM 16-h PP. CWFT. Shoot tips from in vitro seedlings produced 4–5 cm shoots with Lakshmi Sita et al. (1992)
BA ? 0.46 or 4.6 lM Kin 1200 lx. 25 C. 4–5 nodes. Shoot tips from in vitro grown shoots produced up to
(SIM). MS ? RH NR 8 shoots 3–5 cm long on B5 ? 4.44 lM BA ? 4.65 lM Kin
5.71–28.59 lM IAA or within 4–6 w. Nodal explants produced 1 shoot/explant. 80%
4.9–24.42 lM IBA rooting on IAA vs. 30–40% on IBA in RIM. 5.71–11.43 lM
(RIM). pH 5.6–5.8. 2% IAA produced adventitious roots but 28.59 lM IAA produced a
sucrose. 0.8% agar. thick tap root. Acclimatization in sterilized soil ? sand (1:1)
Subculture every 4–5 w with 50% survival. 60% survival when kept in liquid RIM for 2
w then transferred to test tubes with only water for 1 w and the
finally to plastic covered pots
‘B5 ? 0.05% AC ? NR 40–70% germination in 24 h. Only shoot tips, nodes and Anuradha and Pullaiah (1999a)
0.44 lM BA (SG). B5 ? mesocotyls formed shoot buds and multiple shoots; the five other
8.88 lM BA (SIM). ‘MS explants induced callus. Mesocotyls formed maximum
? 0.57 lM IAA ? shoots/explant (7–8). MS and WPM not as effective as B5 as
0.49 lM IBA ? 0.53 lM SIM basal medium. Acclimatization claimed in sand ? soil, but
NAA (RIM). pH 5.7–5.8. not quantified
3% sucrose. Gelling agent
NR
MS ? 2% sucrose ? 0.6% 16-h PP. Light 10.4 shoots/cotyledon on SIM. No shoots formed from epicotyls, Arockiasamy et al. (2000)
agar (SG). MS ? source NR. hypocotyls or internodes. 76.2% of cultures rooted.
0.53 lM NAA ? 1200 lx. 26 C. Acclimatization claimed in sterilized garden soil ? sand (1:1)
4.44 lM BA ? 4.65 lM RH NR and watered with RIM for 1 w, but not quantified
Kin (SIM). MS ?
5.71 lM IAA (RIM).
Carbon source, gelling
agent, pH NR
MS, WPM, Anderson or Chaturani et al. (2006)
Vitis basal medium ? AC 12-h PP. Light 90% germination after 15 min exposure to 0.1% HgCl2. \3 cm
conc. NR or without AC source NR. 23 ± pods were either seedless or with fragile seeds. About 95% of
(SG). pH NR. 3% sucrose. 2 C. RH NR seeds from pods [5 cm in size germinated. No correlation
08% agar between size and germination period. 96% germination in pods
stored for 1 w at 28 ± 5 C. Storage period and browning were
inversely proportional to germination efficiency. Rapid (within 6
d) germination (92%) with 10 mm hypocotyls was possible from

J. A. Teixeira da Silva et al.

seeds cultured on Anderson medium without AC.
Liquid MS ? 4.4 lM BA ? Subculture on paper bridges improved nodal explant survival more Prakash et al. (2006)
2.2 lM TDZ (SIM, 16-h PP. CWFT. 50 than all antioxidants tested but 80% of cultures showed
SMM) with 6-w lE m-2 s-1. 25 ± browning. 74–75% of nodes formed axillary shoots. An 8.3-fold
subcultures on paper 2 C. 65% RH increase in shoots by the 6th subculture. 70% survival after
bridges. ‘MS ? 4.9 lM acclimatization in autoclaved soil ? FYM (4:1) after 5 months
IBA (RIM). pH 5.8. 3%
sucrose. 0.8% agar (RIM
only)
 Red sandalwood (Pterocarpus santalinus L. f.): biology, importance, propagation and…
Table 2 continued
Culture medium, PGRs, Culture conditionsa Experimental outcome, maximum productivity, acclimatization References
additives, subcultures and variation

MS ? 13.32 lM BA (SIM, 17 shoots/seed explant on SIM, then SMM, with 90% explant Padmalatha and Prasad (2008)
nodes). MS ? 4.44 lM 16-h PP. CWFT. response. Rooting data NR. Low (20%) survival of acclimatized
BA ? 9.30 lM Kin (SIM, 83.6 lE m-2 s-1. 25 plantlets in Soilrite ? manure ? sand (1:1:1)
seeds). MS ? 4.44 lM ± 2 C. 60–70% RH
BA (SMM, nodes, seeds).
Auxin-free MS ? 0.25%
phytagel (RIM). pH NR.
3% sucrose. 0.6% agar
MS ? 2.5 lM BA ? 2 lM Max. of 4/4 shoots/cotyledonary node in 2nd subculture in 95% of Rajeswari and Paliwal (2008)
2iP (SIM, SMM). Dip in 16-h PP. CWFT. cotyledonary nodes. 82.5% of shoots induced roots. 95%
5 lM IAA ? 1 lM IBA 60 lmol m-2 s-1. survival of acclimatized plantlets in coarse sand ? clay ? FYM
(ex vitro RIM). pH NR. 24 ± 2 C. RH NR (1:1:1). Tissue-cultured plants showed better morphological
3% sucrose. 0.8% performance than seedlings
Bactoagar
MS ? 1 mg/l BA ? % SG NR. 83% of shoot tips formed new shoots, with 11 Balaraju et al. (2011)
0.45 lM TDZ (SIM). 16-h PP. CWFT. buds/explant after 45 d. 60% of shoots rooted. 73.3% survival of
Subculture every 4 w. MS 35–50 lmol m-2 acclimatized plantlets in organic manure and garden soil ? sand
? 2.22 lM BA ? s-1. 22 ± 1 C. RH (1:1) under in vitro culture conditions. RAPD used to confirm
0.28 lM GA3 (shoot NR lack of variation
elongation). MS ?
0.49 lM IBA (RIM)
using 3–4 cm long shoots
with 4–5 leaves. After 4
w, transfer to PGR-free
MS (root elongation). pH
5.8. 2% sucrose. 0.8%
agar
MS ? 2 lM GA3 (SG). MS 85% of adventitious shoots elongated. 8.8 shoots/shoot tip. 85% Vipranarayana et al. (2012)
? 1 mg/l BA ? 0.5 mg/l 16-h PP. CWFT. rooting after 4 w in soil ? manure (1:1) (survival NR)
NAA (SIM). Pulse in 50 lmol m-2 s-1.
1.5 g/l IBA ? ‘MS 25 ± 2 C. 65% RH
(RIM). pH 5.8. 3%
sucrose. 0.8% agar
MS, WPM or B5 ? 80% survival from immature cuttings when surface sterilized with Warakagoda and Subasinghe (2013)
4–12 lM BA ? 16-h PP. CWFT. 15% Clorox for 10 min. 0.1% AC with WPM was best
0.5–2 lM NAA (SIM). 1220 lx. 23 ± 2 C. interaction. No significant difference in interaction between
25–2500 mg/l IBA pulse 60% RH media and explant type. Maximum number of shoot buds
treatment 12 h ? ‘ MS (- 4.95)/cotyledonary nodal explant on B5 medium
? 0.5 lM IBA (RIM). pH supplemented with 8 lM BA and 2 lM NAA. Longest roots in
5.8. 3% sucrose. 0.01% 25 mg/L pulse treatment. Ex vitro rooting using 1000 mg/L IBA
123

myo-inositol. Gelling produced 40% rooting. Acclimatization performed in sand ?

agent NR coir dust (1:1) was 80% when kept in humid conditions and weak
light for the first 4 w then for 6 w in greenhouse

751
752 J. A. Teixeira da Silva et al.

regulator, PP photoperiod, RAPD random amplified polymorphic DNA, RH relative humidity, RIM root induction medium, SG seed germination, SIM shoot induction medium, SMM shoot
AC activated charcoal, Anderson medium (Anderson 1980), B5 medium, or Gamborg medium (Gamborg et al. 1968), BA N6-benzyladenine (BA is used throughout even though BAP (6-

The original light intensity reported in each study has been represented since the conversion of lux to lmol m-2 s-1 is different for different illumination (main ones represented): for
benzylamino purine) may have been used in the original (Teixeira da Silva 2012), CWFT white fluorescent tubes, d day(s), FYM farmyard manure, GA3 gibberellic acid, IAA indole-3-acetic
acid, IBA indole-3-butyric acid, Kin kinetin (6-furfuryl aminopurine), MS Murashige and Skoog, (1962) medium, NAA a-naphthaleneacetic acid, NR not reported in the study, PGR plant growth
Conclusions and future perspectives

multiplication medium, TDZ thidiazuron (N-phenyl-N’- 1,2,3-thiadiazol-5-ylurea), w week(s), Vitis medium (Chee and Pool 1987), WPM woody plant medium (Lloyd and McCown 1980)
This review highlights key advances in the tissue culture-
Ashrafee et al. (2014) based biotechnology of economically important Pterocar-
pus santalinus. To date, effective protocols for seed surface
disinfection and in vitro germination exist. There are also
effective protocols for direct shoot regeneration from a
References

range of explants or through callus induction. In most

fluorescent lamps, 1 lmol m-2 s-1 = 80 lx; the sun, 1 lmol m-2 s-1 = 55.6 lx; high voltage sodium lamp, 1 lmol m-2 s-1 = 71.4 lx (Thimijan and Heins 1983)
cases, explants are derived from seeds or seedlings which
are not suitable for clonal propagation (Table 1). There-
fore, a clonal method should be developed from vegetative
tissues of elite germplasm. Rooting and survival of
micropropagated plants remain a major limitation to the
success of P. santalinus tissue culture and should be opti-
mized in the future, for example by using CO2 enrichment
pathogenic bacterial isolates of Aeromonas and Pseudomonas
Experimental outcome, maximum productivity, acclimatization

82.1% of leaves and 100% of internodes formed callus. Paste

and vessels that allow for maximized aeration without

made from callus used for antifungal assays against 10

impacting relative humidity levels within the culture vessel

(Teixeira da Silva et al. 2005). The ability to stably pro-
duce units that allow for germplasm conservation would
then stimulate the need for cryoconservation (Teixeira da
Silva and Engelmann 2017; Bi et al. 2017), including
through the application of synthetic seeds (Sharma et al.
2013). Analytic hierarchy, which is a multicriteria deci-
sion-making tool, is valuable for incorporating the per-
ceptions of stakeholders when planning the conservation
and restoration of a P. santalinus population (Kukrety et al.
2013a, c). The micropropagation and biotechnology of
another commercially important tree in this genus, P.
marsupium (Indian kino tree), have recently been reviewed
and variation

(Teixeira da Silva et al. 2018).

Acknowledgements The authors thank Dr. Emilia Caboni (Agri-

cultural Research Council (CRA), Fruit Tree Research Centre, Rome,
Italy), Dr. Randall Niedz (U.S. Department of Agriculture, Agricul-
tural Research Service, U.S. Horticultural Research Laboratory, FL,
USA) and Dr. Ivana Gribaudo (Istituto Protezione Sostenibile delle
Piante—CNR, Grugliasco, Italy) for ideas, comments and suggested
improvements to an early version of the manuscript. The authors also
Culture conditionsa

thank Prof. M.N.V. Prasad (Department of Plant Sciences, University

Conditions NR

of Hyderabad, India) for some suggestions on a later version of the

manuscript.

Open Access This article is distributed under the terms of the

Creative Commons Attribution 4.0 International License (http://crea
tivecommons.org/licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided you give
appropriate credit to the original author(s) and the source, provide a
link to the Creative Commons license, and indicate if changes were
made.
Subculture every 2 w. pH,
MS ? 2.26–9.04 lM 2,4-D

2.69 lM NAA (CIM).

Culture medium, PGRs,

carbohydrate source,
additives, subcultures

? 11.1 lM BA ?
Table 2 continued

gelling agent NR

References

Anderson WC (1980) Tissue culture propagation of red and black

raspberries, Rubus idaeus and R. occidentalus. Acta Hort
112(112):13–20
a

123
Red sandalwood (Pterocarpus santalinus L. f.): biology, importance, propagation and… 753

Ankalaiah C, Mastan T, Reddy MS (2017) A study on the density, Gasson P, MacLachlan IR (2010) PCA of CITES listed Pterocarpus
population structure and regeneration of red sanders Pterocarpus santalinus (Leguminosae) wood. IAWA J 31(2):121–138
santalinus (Fabales: Fabaceae) in a protected natural habitat— Gulrajani ML, Bhaumik S, Oppermann W, Hardtmann G (2002)
Sri Lankamalleswara Wildlife Sanctuary, Andhra Pradesh, India. Kinetic and thermodynamic studies on red sandalwood. Indian J
J Threat Taxa 9(9):10669–10674 Fibre Text Res 27(1):91–94
Anuradha M, Pullaiah T (1998) Investigations on germination of IUCN (2018) CAMP workshops on medicinal plants, India (January
Pterocarpus santalinus (red sanders) with special reference to in- 1997) 1998. Pterocarpus santalinus. In: The IUCN red list of
vitro seed culture. Indian For 124:309–314 threatened species. Version 2016.2. http://www.iucnredlist.org/
Anuradha M, Pullaiah T (1999a) Propagation studies of red sanders details/32104/0. Accessed 21 May 2018
(Pterocarpus santalinus L.f.) in vitro—an endangered taxon of Jhansi Rani S, Usha S (2013) Development of RAPD and specific
Andhra Pradesh, India. Taiwania 44(3):311–324 SCAR markers for the identification of Pterocarpus santalinus
Anuradha M, Pullaiah T (1999b) In vitro seed culture and induction of L. J Cell Tissue Res 13:3809–3816
enhanced axillary branching in Pterocarpus santalinus and a Jyothi CP, Chandrashekar R, Lakshmi B (2014) Isolation of
method for rapid multiplication. Phytomorphology 49:157–163 Pterocarpus santalinus L. genomic DNA, for quality check
Arockiasamy S, Ignacimuthu S, Melchias G (2000) Influence of and quantification with reference to Telangana region, Andhra
growth regulators and explant type on in vitro shoot propagation Pradesh, India. Indian J Sci 8:21–24
and rooting of red sandal wood (Pterocarpus santalinus L.). Kalimuthu K, Lakshmanan KK (1995) Effect of different treatments
Indian J Exp Biol 38(12):1270–1273 on pod germination of Pterocarpus species. Indian J For
Arunkumar AN, Joshi G (2014) Pterocarpus santalinus (red sanders) 18:104–106
an endemic, endangered tree of India: current status, improve- Kaner J, Jiufang L, Yongji X, Ioraş F (2013) A re-evaluation of
ment and the future. J Trop For Environ 4:1–10 woods used in Chinese historic furniture (part two). Bull
Arunkumara KKIU, Walpola BC, Subasinghe S, Yoon M-H (2011) Transilv Univ Brasov Ser II For Wood Ind Agric Food Eng
Pterocarpus santalinus Linn. f. (Rath handun): a review of its 6:31–40
botany, uses, phytochemistry and pharmacology. J Korean Soc Kedharnath S, Rawat MS, Uniyal DP, Kantham DL (1976) Studies on
Appl Biol Chem 54:495–500 field grafting and the growth of grafts in red sanders. Indian For
Ashrafee TS, Rahman MM, Chakraborty A, Prodhan SH (2014) 102:761–765
Antibacterial potentiality of red sandalwood callus against Kesava Reddy K, Srivasuki KP (1990) Vegetative propagation of red
pathogenic isolates of Aeromonas and Pseudomonas. Univers J sanders (Pterocarpus santalinus Linn.). Indian For 116:536–540
Plant Sci 2(4):86–91 Kukrety S, Dwivedi P, Jose S, Alavalapati JRR (2013a) Stakeholders’
Azamthulla M, Balasubramanian R, Kavimani S (2015) A review on perceptions on developing sustainable Red Sanders (Pterocarpus
Pterocarpus santalinus Linn. World J Pharm Res 4:282–292 santalinus L.) wood trade in Andhra Pradesh, India. For Policy
Balaraju K, Agastian P, Ignacimuthu S, Park K (2011) A rapid in vitro Econ 26(C):43–53
propagation of red sanders (Pterocarpus santalinus L.) using Kukrety S, Gezan S, Jose S, Alavalapati JRR (2013b) Facilitating
shoot tip explants. Acta Physiol Plant 33(6):2501–2510 establishment of advance regeneration of Pterocarpus santalinus
Banerjee A, Mukherjee AK (1981) Chemical aspects of santalin as a L.—an endangered tree species from India. Restor Ecol
histological stain. Stain Technol 56(2):83–85 21(3):372–379
Berliner N (1996) Beyond the screen: Chinese furniture of the 16th Kukrety S, Jose S, Alavalapati JRR (2013c) Exploring stakeholders’
and 17th centuries. Mus Fine Arts, Boston perceptions with analytic hierarchy process—a case study of red
Bi W-L, Pan C, Hao X-Y, Cui Z-H, Kher MM, Marković Z, Wang sanders (Pterocarpus santalinus L.) restoration in India. Restor
Q-C, Teixeira da Silva JA (2017) Cryopreservation of grapevine Ecol 21(6):777–784
(Vitis spp.)—a review. In Vitro Cellular Dev Biol Plant Kumar A, Gopal M (1975) A note on temperature sensitivity for
53(5):449–460 germination of Pterocarpus santalinus seed. Seed Sci Technol
Botanical Survey of India (2012) Identification protocol for red 3:605–613
sanders. In: Pharmacognosy of negative listed plants. Ministry of Kumar N, Ravindranath B, Seshadri TR (1974) Terpenoids of
Environment and Forests, India, pp 171–181. http://wccb.gov.in/ Pterocarpus santalinus heartwood. Phytochemistry 13:633–636
WriteReadData/userfiles/file/PROTOCOL%20FOR%20Red% Kumarasinghe HKMS, Subasinghe S, Arunakumara KKIU (2003)
20sanders%20identification.pdf Studies on seed propagation of rathhadun (Pterocarpus santali-
Chaturani GDG, Jayatilleke MP, Subasinghe S (2005) In vitro callus nus Linn.) as affected by method of soaking and scarification. In:
formation of red sandalwood (Pterocarpus santalinus L.) as Proceedings of 9th annual forestry and enviorenmcnt sympo-
affected by explant type and different levels of 2,4-D and BAP. sium, p 57
In: Proceedings of 10th annual forestry & environmental Lakshmi Sita G, Sreenatha KS, Sujata S (1992) Plantlet production
symposium. University of Sri Jayewardenpura, Gangodawila, from shoot tip cultures of red sandalwood (Pterocarpus santali-
Sri Lanka, p 37 nus L.). Curr Sci 62:532–535
Chaturani GDG, Subasinghe S, Jayatilleke MP (2006) In-vitro Leifert C, Morris CE, Waites WM (1994) Ecology of microbial
establishment, germination and growth performance of red saprophytes and pathogens in tissue culture and field-grown
sandalwood (Pterocarpus santalinus L.). Trop Agric Res Ext plants: reasons for contamination problems in vitro. Crit Rev
9:116–130 Plant Sci 13:139–183
Chee R, Pool RM (1987) Improved inorganic media constituents for Lloyd G, McCown B (1980) Commercially-feasible micropropaga-
in-vitro shoot multiplication of Vitis. Sci Hortic 32:85–89 tion of mountain laurel, Kalmia latifolia, by use of shoot-tip
Dayanand T, Lohidas T (1988) Effect of different treatments on pod culture. Int Plant Propag Soc Proc 30:421–427
germination of red sanders (Pterocarpus santalinus Linn. f). Murashige T, Skoog F (1962) A revised medium for rapid growth and
Indian J For 11:87–88 bioassays with tobacco tissue cultures. Physiol Plant 15:473–497
Gamborg OLL, Miller RAA, Ojima K (1968) Nutrient requirements Naidu CV (2001a) Improvement of seed germination in red sanders
of suspension cultures of soybean root cells. Exp Cell Res (Pterocarpus santalinus Linn. F.) by plant growth regulators.
50(1):151–158 Indian J Plant Physiol 6:205–207

123
754 J. A. Teixeira da Silva et al.

Naidu CV (2001b) Seed pretreatment methods to improve germina- sanders (Pterocarpus santalinus). In: Pandey K, Ramakantha V,
tion red sanders (Pterocarpus santalinus) Linn. F. Indian J For Chauhan S, Arun Kumar A (eds) Wood is good. Springer
24:342–343 Singapore, Singapore, pp 139–151
Naidu CV, Rajendrudu G (2001) Influence of kinetin and nitrogenous Teixeira da Silva JA (2012) Is BA (6-benzyladenine) BAP (6-
salts on seed germination of red sanders (Pterocarpus santalinus benzylaminopurine)? Asian Aust J Plant Sci Biotechnol 6(Spe-
Linn. f.). Seed Sci Technol 29:669–672 cial Issue 1):121–124
Navada KK, Vittal RR (2014) Ethnomedicinal value of Pterocarpus Teixeira da Silva JA, Engelmann F (2017) Cryopreservation of oil
santalinus (Linn. f.), a Fabaceae member. Orient Pharm Exp palm (Elaeis guineensis Jacq.). Cryobiology 77:82–88
Med 14:313–317 Teixeira da Silva JA, Giang DTT, Tanaka M (2005) Micropropaga-
Padmalatha K, Prasad MNV (2007) Morphological and molecular tion of sweet potato (Ipomoea batatas) in a novel CO2-enriched
diversity in Pterocarpus santalinus L. f—an endemic and vessel. J Plant Biotechnol 7(1):67–74
endangered medicinal plant. Med Aromat Plant Sci Biotechnol Teixeira da Silva JA, Kher MM, Soner D, Nataraj M (2016a) African
1:263–273 sandalwood or Nepalese sandalwood: a brief synthesis. Notulae
Padmalatha K, Prasad MNV (2008) In vitro plant regeneration of Scientia Biologicae 8:57–61
Pterocarpus santalinus L.f (red sanders)—an endangered medic- Teixeira da Silva JA, Kher MM, Soner D, Page T, Zhang X, Nataraj
inal plant and important timber tree. Tree For Sci Biotechnol M, Ma G (2016b) Sandalwood: basic biology, tissue culture, and
2:1–6 genetic transformation. Planta 243:847–887
Patel HS, Tandel MB, Prajapati VM, Amlani MH, Prajapati DH Teixeira da Silva JA, Kulus D, Zhang X, Zeng SJ, Ma GH, Piqueras A
(2018) Effect of different pre-sowing treatments on germination (2016c) Disinfection of explants for saffron (Crocus sativus L.)
of red sanders (Pterocarpus santalinus L. f.) in net house tissue culture. Environ Exp Biol 14(4):183–198
condition. Int J Chem Stud 6:876–879 Teixeira da Silva JA, Kher MM, Soner D, Nataraj M, Dobránszki J,
Prakash E, Sha Valli Khan PS, Sreenivasa Rao TJV, Meru ES (2006) Millar MA (2017a) Santalum molecular biology: molecular
Micropropagation of red sanders (Pterocarpus santalinus L.) markers for genetic diversity, phylogenetics and taxonomy, and
using mature nodal explants. J For Res 11:329–335 genetic transformation. Agrofor Syst. https://doi.org/10.1007/
Rajeswari V, Paliwal K (2008) In vitro plant regeneration of red s10457-017-0075-8 (in press)
sanders (Pterocarpus santalinus L.f.) from cotyledonary nodes. Teixeira da Silva JA, Sharma M, Hossain M, Dobránszki J, Cardoso
Indian J Biotechnol 7:541–546 JC, Zeng SJ (2017b) Acclimatization of in vitro-derived
Raju KK, Nagaraju A (1999) Geobotany of red sanders (Pterocarpus Dendrobium. Hortic Plant J 3(3):110–124
santalinus)—a case study from the southeastern portion of Teixeira da Silva JA, Kher MM, Soner D, Nataraj M (2018) Indian
Andhra Pradesh. Environ Geol 37:340–344 kino tree (Pterocarpus marsupium): propagation, micropropaga-
Ramabrahmam V, Sujatha M (2016) Red sanders in Rayalaseema tion, and biotechnology. Environ Exp Biol 16(1):1–8
region of Andhra Pradesh: importance to commercial & Thimijan RW, Heins RD (1983) Photometric, radiometric, and
medicinal value. IOSR J Pharm Biol Sci 11:57–60 quantum light units of measure: a review of procedures for
Ramakrishna A (1962) The red sanders and its future. Indian For interconversion. HortScience 18:818–822
88:202–206 Usha R, Rani SJ, Prasuna TG (2013) Genetic relationship between
Rao SP, Raju AJS (2002) Pollination ecology of the red sanders quality and non quality wood of Pterocarpus santalinus L. (red
Pterocarpus santalinus (Fabaceae), an endemic and endangered sanders), an endemic tree species by using molecular markers.
tree species. Curr Sci 83:1144–1148 J Chem Pharm Sci 6:189–194
Rao SP, Atluri JB, Reddi CS (2001) Intermittent mass blooming, Vedavathy S (2004) Cultivation of endemic red sanders for interna-
midnight anthesis and rockbee pollination in Pterocarpus tional trade. Nat Prod Radiance 3:83–84
santalinus (Fabaceae). Nord J Bot 21:271–276 Vijayalakshmi KP, Renganayaki PR (2017) Preliminary study on
Rawat MS, Uniyal DP (1996) Identification of wavy grained red germination of pre-treated seed of red sanders under nursery
sanders (Pterocarpus santalinus) at nursery stage. Indian For conditions from Tamil Nadu. Agric Update 12:956–959
122:831–833 Vipranarayana S, Prasad TNVKV, Damodharam T (2012) In vitro
Ruffoni B, Savona M (2013) Physiological and biochemical analysis seed germination and induction of enhanced shoot multiplication
of growth abnormalities associated with plant tissue culture. in Pterocarpus santalinus Linn. f: an endemic medicinal plant of
Hortic Environ Biotechnol 54:191–205 Seshachalam Hills, Tirumala. Int J Pure Appl Sci Technol
Sarita S, Bhatnagar SP, Bhojwani SS (1988) Preliminary investiga- 9:118–126
tions on micropropagation of leguminous timber tree Pterocar- Warakagoda PS, Subasinghe S (2013) In vitro propagation of
pus santalinus. Phytomorphology 38:41–45 Pterocarpus santalinus L. (red sandalwood) through tissue
Sarma CR (1993) Study of outturn and value of red sanders. J Trop culture. J Natl Sci Found Sri Lanka 41:53–63
For 9:125–131 Wu SF, Chang FR, Wang SY, Hwang TL, Lee CL, Chen SL, Wu CC,
Sen Gupta PC, Mukherjee AK (1981) Newer applications of the Wu YC (2011) Anti-inflammatory and cytotoxic neoflavonoids
histological stain prepared from Pterocarpus santalinus. Stain and benzofurans from Pterocarpus santalinus. J Nat Prod
Technol 56:79–82 74:989–996
Sharma S, Shahzad A, Teixeira da Silva JA (2013) Synseed Zhang F, Xu C-H, Li M-Y, Huang A-M, Sun S (2014) Rapid
technology—a complete synthesis. Biotechnol Adv 31:186–207 identification of Pterocarpus santalinus and Dalbergia louvelii
Suresh K, Hegde M, Deenathayalan P, Karthick Kumar P, Thanga- by FTIR and 2D correlation IR spectroscopy. J Mol Struct
pandi M, Gurudev Singh B, Krishnakumar N (2017) Variation in 1069:89–95
heartwood formation and wood density in plantation-grown red

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