Beruflich Dokumente
Kultur Dokumente
net/publication/283132113
CITATIONS READS
10 22,512
4 authors, including:
Some of the authors of this publication are also working on these related projects:
Development of sustainable weed management practices in diversified cropping systems View project
All content following this page was uploaded by Saurabh Pagare on 11 August 2016.
of the cell (organism) with its environment. These cell specific and these are low molecular weight
compounds are often involved in plants protection compounds. These compounds often differ
Members Copy, Not for Commercial Sale
against biotic or abiotic stresses. Secondary between individuals from the same population
www.IndianJournals.com
metabolites are from different metabolites of plants in respect of their amount and types.
families that can be highly inducible in response They protect plants against stresses, both biotic
to stresses. Primary metabolites perform (bacteria, fungi, nematodes, insects or grazing
essential metabolic roles by participating in by animals) and abiotic (higher temperature and
nutrition and reproduction. A few SMs are used moisture, shading, injury or presence of heavy
as especially chemical such as drugs, flavours, metals). SMs are used as especially chemical
fragrances, insecticides, and dyes and thus have such as drugs, flavours, fragrances, insecticides,
a great economic value. These new technologies and dyes by human because of a great economic
will serve to extend and enhance the continued value.
usefulness of the higher plants as renewal
In plants, SMs can be separated into three
sources of chemicals, especially medicinal
groups (Terpenoids, Polyketides and
compounds. A continuation and intensification
Phenypropanoids) based on their biosynthesis
efforts in this field is expected to lead to
origin (1). Alkaloids are additional class of SMs,
successful biotechnological production of
which are nitrogenous organic molecules
specific, valuable and as yet unknown plant
biosynthesized mainly from amino-acids, e.g.,
chemicals.
tryptophan, tyrosine, phenylalanine, lysine and
Keywords: Secondary metabolites, drugs, arginine using many unique enzymes (2). Many
flavours, fragrances, biotechnology of the most important therapeutic agents are
alkaloids. The sites of biosynthesis are
Introduction compartmentalised at cellular or sub-cellular
Plants possess capacity to synthesize level. However SMs can be transported long
different organic molecules called secondary distances and accumulate from their location of
metabolites. Unique carbon skeleton structures synthesis.
are basic properties of plant secondary Primary Vs Secondary Metabolites : Primary
metabolites. Secondary metabolites are not metabolites are found in all plants and execute
necessary for a cell (organism) to live, but play a vital metabolic responsibilities, by participating in
nutrition and reproduction (2). Sometimes it is beetle, wasps, moths, bees, etc and a popular
hard to discriminate primary and secondary ingredient in commercial insecticides because
metabolites. For example, both primary and of low persistence in the environment and low
secondary metabolites are found among the mammalian toxicity. In Gymnoperms (conifers)
terpenoids and the same compound may have á-pinene, â-pinene, limonene and myrecene are
both primary and secondary roles. Secondary found. A number of sesquiterpenes have been
metabolites are broad range of compounds from till now reported for their role in plant defense
different metabolite families that can be highly such as costunolides are antiherbivore agents
inducible in stress conditions. Carotenoids and of family composite characterized by a five
flavonoids are also involved in cell pigmentation member lactone rings (a cyclic ester) and have
in flower and seed, which attract pollinators and strong feeding repellence to many herbivorous,
seed dispersers. Therefore, they are also insects and mammals. ABA is also a
involved in plant reproduction (3). Plant primary sesquiterpene plays primarily regulatory roles in
products refer to the compounds of nucleic acids, the initation and maintenance of seed and bud
proteins, carbohydrates, fats and lipids and are dormancy and plants response to water stress
related to structure, physiology and genetics, by modifying the membrane properties and act
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
which imply their crucial role in plant as a transcriptional activator (4). Abietic acid is a
development. In contrast, secondary metabolites diterpene found in pines and leguminous tress.
It is present in or along with resins in resin canals
Members Copy, Not for Commercial Sale
concentrations. Primary metabolism refers to the of the tree trunk . Another compound phorbol
processes producing the carboxylic acids of the (Diterpene ester), found in plants of
Krebs cycle. Secondary metabolites, on the other euphorbiaceae and work as skin irritants and
hand, are non-essential to life but contribute to internal toxins to mammals. The milkweeds
the species’ fitness for survival. In fact, the produce several better tasting glucosides
specific constituents in a certain species have (sterols) that protect them against herbivors by
been used to help with systematic determination, most insects and even cattle. Several high
groups of secondary metabolites being used as molecular weight polyterpenes occur in plants.
markers for botanical classification The principal tetraterpenes are carotenoids
(chemotaxonomy). Plants secondary metabolites family of pigments.
can be divided into three chemically distinct
groups viz: Terpenes, Phenolics, N (Nitrogen) and (II) Phenolic compounds : Plants produce a
S (sulphur) containing compounds. large variety of secondary products that contain
a phenol group, a hydroxyl functional group on
I)Terpenes : Terpenes comprise the biggest an aromatic ring called Phenol, a chemically
group of secondary metabolites and are free by heterogeneous group also. They could be an
their common biosynthetic origin from acetyl-coA important part of the plants defence system
or glycolytic intermediates. An immense bulk of against pests and disease including root parasitic
the diverse terpenes structures produced by nematodes (5). Elevated ozone (mean 32.4ppb)
plants as secondary metabolites that are increased the total phenolic content of leaves and
supposed to be concerned in defense as toxins had minor effects on the concentration of
and feeding deterrents to a large number of plant individual compounds (6).Coumarin are simple
feeding insects and mammals. Terpenes are phenolic compounds widespread in vascular
divided into monoterpenes, sesquiterpenes, plants and appear to function in different
diterpene, Triterpenes and polyterpenes. The capacities in various plant defense mechanisms
pyrethroid (monoterpenes esters) occur in the against insect herbivores and fungi. They derived
leaves and flowers of Chrysanthemum species from the shikimic acid pathway, common in
show strong insecticidal responses to insects like bacteria, fungi and plants but absent in animals
Saurabh Pagare et al
Current Trends in Biotechnology and Pharmacy 296
Vol. 9 (3) 293-304 July 2015, ISSN 0973-8916 (Print), 2230-7303 (Online)
(7). Some coumarin derivatives have higher anti- were the most active enzymes in red cabbage
fungal activity against a range of soil borne plant seedlings exposed to Cu++ stress (11).Tannins
pathogenic fungi and exhibit more stability as included in the second category of plant phenolic
compared to the original coumarin compounds polymers with defensive properties. Tannins are
alone (7). Furano is Also a type of coumarin with general toxins that significantly reduce the growth
special interest of phytotoxicity, abundant in and survivorship of many herbivores, and also
members of the family umbelliferae including act as feeding repellents to a great diversity of
celery parsnip and parsley. Psoraline, basic linear animals.
furacoumarin, known for its use in the treatment
(III) Sulphur containing secondary
of fungal defence and found very rarely in SO2
metabolites: They include GSH, GSL,
treated plants (8). Ligin is a highly branched
Phytoalexins, Thionins, defensins and allinin
polymer of phenyl- propanoid groups, formed
which have been linked directly or indirectly with
from three different alcohols viz., coniferyl,
the defence of plants against microbial pathogens
coumaryl and synapyl which oxidized to free
(12,13,14). GSH is the one of the major form of
radical (ROS) by a ubiquitous plant enzyme-
organic sulphur in the soluble fraction of plants
peroxidises, reacts simultaneously and randomly
and has an important role as a mobile tool of
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
invading pathogens by accumulating around the monocots and gymnosperms. Generally, most of
site of infection, appears to a common them, including the pyrrolizidine alkaloids (PAs)
mechanism of resistance to pathogenic microbes are toxic to some degree and appear to serve
in a wide range of plants. Many of these changes primarily in defense against microbial infection
are linked to a rapid apoptotic response, resulting and herivoral attack. Cyanogenic glucosides
in death of one or a few invaded plant cells, constitue a group of N-containing protective
known as the hypersensitive response (HR). compounds other than alkaloids, release the
Most plant families produce organic phytolexins poison HCN and usually occur in members of
of diverse chemistry; these groups are often families viz., Graminae, Roosaceae and
associated with a family, for example leguminosesae. They are not themselves toxic
sesquiterpenoids of Solanaceae, isoflavonoids but are readily broken down to give off volatile
of Leguminosae, while phytoalexins from poisonous substance like HCN and volatile H2S
Brassica have an indole or related ring system when the plant crushed; their presence deters
and one S atom as common structural features. feeding by insects and other herbivorous such
Crucifereae appears to be the only plant family as snails and slugs. Amygdalin, the common
producing these S metabolites, which are clearly cynogenic glucoside found in the seeds of
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
different from the other well- known almonds, apricot, cherries and peaches while
GSL.Cruciferous crops are cultivated worldwide Dhurrin, found in Sorghum bicolar.
because they are extremely valuable and for the Many plants also contain unusual amino acids
Members Copy, Not for Commercial Sale
www.IndianJournals.com
last decades, various research groups have called non-protein amino-acids that incorporated
investigated cruciferous phytoalexins as well as into proteins but are present as free forms and
their biological activity.Typically, there are multiple act as protective defensive substance. For
responses involving several related derivatives examples, canavanine and azetidine-2 carboxylic
such as up to nine wyerone (Furano-acetylenic acid are close analogs of arginine and proline
derivatives) forms in Vicia fava and several forms respectively. They exert their toxicity in various
of phaseollin in Phaseolus vulgaris and glyceollin ways. Some block the synthesis of or uptake of
in Glycine max, postin in Pisum sativum pods, protein amino acid while others can be mistakenly
Ipomearone in sweet pototo, orchinol in orchid incorporated into proteins. Plants that
tubers, trifolirhizin in red clover. Defensins, synthesized non-protein amino acid are not
thionins and lectins are S-rich non-storage plant susceptible to the toxicity of these compounds
proteins synthesize and accumulate after but gain defence to herbivorous animals, insects
microbial attack and such related situations. They and pathogenic microbes.
inhibit growth of a broad range of fungi.
Additionally defensins genes are partly pathogen- Transport, Storage and Turnover: SMs can
inducible and others that are involved in be water soluble (hydrophilic) compounds or
resistance can be expressed constitutively. Some lipophilic (needs organic solvents), therefore
plant species produce lectins as defensive needs different cellular mechanism for their
proteins that bind to carbohydrates or transport, storage and turnover. Most substances
carbohydres containing proteins. are synthesized in the cytoplasm, the ER or in
the organelles. Hydrophilic SMs are usually
(IV) Nitrogen containing secondary stored in the vacuole after their formation in
metabolites: They include alkaloids, cyanogenic cytoplasm, whereas lipophilic substances are
glucosides, and non-proteins amino-acids. Most sequestered in resin ducts, laticifers, glandular
of them are biosynthesized from common amino- hairs, trichomes, thylakoid membranes or on the
acids. Alkaloids found in approximately 20% of cuticle. Hydrophilic SMs have to pass the
the species of vascular plants, most frequently tonoplast, which is impermeable to many of the
in the herbaceous dicot and relatively a few in polar secondary metabolites. For some alkaloids
Saurabh Pagare et al
Current Trends in Biotechnology and Pharmacy 298
Vol. 9 (3) 293-304 July 2015, ISSN 0973-8916 (Print), 2230-7303 (Online)
and flavanoids, a specific transporter has been extraction process. Successful extraction begins
described, which pumps the compounds into the with careful selection and preparation of plant
vacuole. In order to avoid autotoxicity, plants samples. During the extraction of plant material,
cannot store these compounds in the vacuole it is important to minimize interference from
but usually sequester them on the cuticle, in dead compounds that may coextract with the target
resin ducts or cells which are lined by a compounds, and to avoid contamination of the
biomembrane but an impermeable solid barrier. extract, as well as to prevent decomposition of
In many instances, the site of biosynthesis is important metabolites or artifact formation as a
restricted to a single organ such as roots, leaves result of extraction conditions or solvent
or fruits, but an accumulation of the impurities. Researchers from a variety of
corresponding products can be detected in scientific disciplines are confronted with the
several other plant tissues. Long distance challenge of extracting plant material with
transport must take place in these instances. The solvents, often as a first step toward isolating and
xylem or phloems are likely transport routes but identifying the speciûc compounds responsible
an apoplastic transport can also be involved. for biological activities associated with a plant
Storage can also be tissue and cell- specific, or a plant extract. The impetus for this research
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
depending upon the protection providing to the arises largely because plants form the foundation
plants.In a number of plants, specific idioblasts of traditional pharmacopeias, and because many
have been detected that contain tannins, of our currently important pharmaceutical drugs
Members Copy, Not for Commercial Sale
www.IndianJournals.com
alkaloids or glucosinolates. More often, SMs are are obtained from plants. Further interest arises
concentrated in trichomes or glandular hairs from the growing awareness that many of the
(many terpenoids in Labiatae, Asteraceae), secondary metabolites of organisms, including
stinging hairs (many amines in urticaceae) or the plants, serve important biological and ecological
epidermis itself (many alkaloids, flavanoids, roles, mainly as chemical messengers and
anthocyanins, cynogenic glycosides, coumarins, defensive compounds. Investigators engaged in
etc.) flowers, fruits and seeds are usually rich in the isolation of secondary metabolites from plants
SMs, especially in annual plants. In perennial soon discover the need for considerable
species, high amounts of SMs found in bulbs, laboratory finesse in the apparently routine
roots, rhizomes and the bark of roots and stems. ‘‘sample preparation’’ steps that convert crude
It is well-established that profiles of SMs vary with plant material into an extract suitable for chemical
time, space and developmental stage. Since analysis, biological testing, or chromatographic
related plant species often show similarities in separation.
the profiles of their SMs, they have been used
as taxonomic tool in plant systematic. However, Major Secondary Metabolite Pathways : In
profiles of closely- related plants quite often differ plants particularly three pathways are the source
substantially or those of unrelated plant group of most secondary metabolites: The shikimate
show strong similarities; this clearly shows that pathway, the isoprenoid pathway and the
SM patterns are not unambiguous systematic polyketide pathway. After the formation of the
markers but that convergent evolution and major basic skeletons, further modifications
selective gene expression are common themes. result in plant species specific compounds. The
shikimate pathway is the major source of
Extraction of Secondary Metabolites from aromatic compounds. It is found in
Plant : Plant secondary metabolites are currently microorganisms and plants, but not in mammals,
the subject of much research interest, but their making it an interesting target for herbicides and
extraction as part of phytochemical or biological antibiotics, as these compounds are expected
investigations presents specific challenges that not to have any effect on the mammalian system.
must be addressed throughout the solvent Glyphosate is a well known example. The
enzymes channeling chorismate into the oils and colored flavonoids or tetraterpenes) also
aromatic amino acids pathways are chorismate to attract insects for pollination or other animals
mutase and anthranilate synthase. Although, in for seed dispersion, in this case secondary
several plant species for both chorismate mutase metabolites serve as signal compounds.
and anthranilate synthase more than one gene Compounds belonging to the terpenoids,
has been cloned, only in case of chorismate alkaloids and flavonoids are currently used as
mutase a plastidial and a cytosolic enzyme have drugs or as dietary supplements to cure or
been found. The phenylpropanoid pathway is one prevent various diseases (19) and in particular
of the most important metabolic pathways in some of these compounds seem to be efficient
plants in terms of carbon flux. In a cell more than in preventing and inhibiting various types of
20% of the total metabolism can go through this cancer (20, 21). It has been estimated that 14-
pathway, the enzyme chorismate mutase is an 28% of higher plant species are used medicinally
important regulatory point. The importance of this and that 74% of pharmacologically active plant
pathway is due to the fact that it leads to among derived components were discovered after
others lignin, lignans, flavonoids, and following up on ethno-medicinal use of the plants
anthocyanins. Key to these products is the (22). Secondary metabolites are a metabolic
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
enzyme can be found in all plants, in some plants producing organism and biosynthetized from
a single enzyme is found, whereas others may one or more general metabolites by wider variety
have several iso-enzymes. The other important of pathways than is available in general
pathway in plants is that of the terpenoids, also metabolism.
known as isoprenoid pathway. Terpenoids include Presence of volatile monoterpenes or
more than one third of all known secondary essential oils in the plants provides an important
metabolites. Moreover, the C5-building block is defense strategy to the plants, particularly against
also incorporated in many other skeletons, e.g. herbivorous insect pests and pathogenic fungi.
in anthraquinones, naphtoquinones, These volatile terpenoids also play a vital role in
cannabinoids, furanocoumarines, and terpenoid plant-plant interactions and serve as attractants
indole alkaloids. In the “decoration” type of for pollinators (23). They act as signalling
reactions in various types of secondary molecules and depict evolutionary relationship
metabolites C5-units are attached to the basic with their functional roles. Soluble secondary
skeleton, e.g. hop bitter acids, flavonoids and compounds such as cyanogenic glycosides
isoflavonoids. isoflavoids and alkaloids can also be toxic to
animals.
Functions of Secondary Metabolites : Many
secondary compounds have signalling functions Biotechnology and Secondary Metabolites :
influence the activities of other cells, control their Since SM have evolved as compounds that are
metabolic activities and co-ordinates the important for the ûtness of the organisms
development of the whole plant. Other producing them, many of them interfere with the
substances such as flower colours serve to pharmacological targets, which make them
communicate with pollinators or protect the plants interesting for several biotechnological
from feeding by animals or infections by applications. Controlled clinical studies have
producing specific phytoalexines after fungi shown the efficacy of several, for example
infections that inhibit the spreading of the fungi extracts from Ginkgo biloba, Hypericum
mycelia within the plant (18). Plants use perforatum, Piper methysticum, Chamomilla
secondary metabolites (such as volatile essential recutita, Crataegus monogyna, Silibum
Saurabh Pagare et al
marianum, Melissa ofûcinalis, Mentha piperita, metabolism has already had a direct inûuence
Valeriana ofûcinalis. for example, when Atropa belladonna plants were
transformed with the gene that encodes the
The use of stimulants (such as caffeine,
enzymes converting L-hyoscyamine into L-
nicotine, ephedrine), fragrances (several
scopolamine, new plants were generated which
essential oils), ûavours (essential oils, capsaicin,
produced scopolamine as the major product.
piperine, etc.), natural dyes, poisons (strychnine)
More often, ûavonoid metabolism has been
and hallucinogens (morphine, heroin, cocaine,
altered genetically, producing plants with different
tetrahydro cannabinol) is based on SM. Since
ûower colours. It is a challenge for future
many SM are insecticidal, fungicidal and
research to isolate the genes of biosynthetic
phytotoxic, they may be used in agriculture as
pathways and to express them either in
natural plant protectants. Before the advent of
transgenic plants or in microbes.
synthetic pesticides about 60 years ago, plant-
derived insecticides (including nicotine, rotenone, If successful, recombinant bacteria or
quassin, ryanodine, pyrethrins and azadirachtins) yeasts might be grown someday, which will
were a common theme. Applications produce valuable plant SM. Combinatorial
unequivocally showed that these natural biosynthesis might then be an open field. Using
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
insecticides worked. One ecological advantage genes encoding enzymes for the biosynthesis of
is that SM are readily degraded in plants and in antibiotics, this strategy has already been
Members Copy, Not for Commercial Sale
soil, is also their disadvantage and synthetic successful. It has also brought about renewed
www.IndianJournals.com
pesticides are more resistant and persistent. interest in the regulation of SM synthesis and in
Moreover, modern pesticides are usually more the location and means of sequestration of these
potent than biopesticides. On the other hand, substances within the plant. In recent years,
plants are easy to grow and biopesticides could attempts have been made to express the genes
be a sustainable source of plant protectants for of alkaloid biosynthesis in microorganisms.
farmers in countries that do not have access to Ultimately, it might be possible to produce
Western synthetic pesticides. Unfortunately, valuable alkaloids from recombinant bacteria or
legislation does not favour mixtures of yeast. If the corresponding SM (both from plant
compounds to be used as pesticides; therefore, or microbial origin) confers resistance to insects
the development of biorational pesticides has to or pathogens, genetic transformation of
face many obstacles. Nevertheless, natural susceptible crop plants could be another valuable
compounds do provide an underexplored avenue for Exploitation. For more than two
alternative. As a consequence of these various decades, scientists around the world have tried
applications, a world market for plant extracts to produce valuable SM in cell or organ cultures.
and isolated SM exists, which exceeds 10 billion Whereas undifferentiated cell cultures have often
US dollars annually. Therefore, it is a challenge failed to produce such a compound in reasonable
for biotechnologists to find ways to produce these yields, differentiated organ cultures (e.g.
compounds in sufûcient quantity and quality. transformed root cultures) are often as active as
The main and traditional way is to grow the the intact plant .Cell- and tissue-speciûc gene
respective plants in the ûeld or in greenhouses expression appears to control these processes.
and to extract the products from them. For In addresses the production of SM in vitro (Table
several species, new varieties have been 1).
selected with improved yields and quality. In this It is possible that genetic engineering may
context, cell and organ culture are important help to improve plant cell cultures as
techniques for in vitro propagation. In a few biotechnological production systems in the future.
instances, genetic engineering of secondary
Saurabh Pagare et al
Conclusion response of grape (Vitis vinifera L.)
This review has dealt with a small cv.Malbec leaf Tissues to ultraviolet-B
selection of plant secondary- metabolites and radiation by enhancing ultraviolet –
their potential roles in defence mechanisms and absorbing compounds, antioxidant enzymes
ecological adaptation, in addition to the topics and membrane sterols. Plant cell Environ.
we have covered. there is an enormous range of 33(1):1-10
other compounds present in the plant kingdom,
5. Wuyts, N., De waele, D. and Swennen, R.
with a very varied distribution. Plant secondary
(2006). Extraction and partial characteriza-
metabolism produces products that aid in the
-tion of polyphenol oxidase from banana
growth and development of plants but are not
(Musa acuminate grandr naine) roots. Plant
required for the plant to survive. Secondary
Physiol Biochem. 44:308-314
metabolites have important ecological functions
in plants: They protect plants against being eaten 6. Savirnata, N.M., Jukunen-Titto, R.,
by herbivores and against being infected by Oksanen, E. and Karjalainen, R.O. (2010).
microbial pathogens. They serve as attractants Leaf Phenolic compounds in red clover
(odor, color, taste) for pollinators and seed- (Trfolium Pratense L.) induced by exposure
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
Plant Biologists, Rockville, MD, doi/10.1199/ 25. Lee, S.Y., Xu, H., Kim, Y.K. and Park, S.U.
tab.0009 (2007). Rosmarinic acid production in hairy
17. Lipka, U., Fuchs, R.., Kuhns, C., root cultures of Agastache rugosa Kuntze.
Petutschnig, E. and Lipka, V. (2010). Live World J Microbiol Biotechnol. 20:969-972
and let die-Arabidopsis non-host resistance 26. Krolicka, A., Kartanowicz, R.,Wosinskia, S.,
to powdery mildews. Eur J Cell Biol. Zpitter, A., Kaminski, M. and Lojkowska, E.
89(2):194-199 (2006). Induction of secondary metabolite
production in transformed callus of Ammi
18. Mansfield, J.W. (2000). Antimicrobial
majus L. grown after electromagnetic
compounds and resistances. The role of
treatment of the culture medium. Enz Microb
phytoalexins and phytoanticipins. In:
Technol. 39: 1386 -1389
slusarenko A. J., fraser R.S.S., vanloon L.C.
and fraser R.S. (eds). Mechanism of 27 Staniszewska, I., Krolicka, A., Mali, E.,
resisyance to plant diseases. Springer- Ojkowska, E. and Szafranek, J. (2003).
verlag New York., pp325-363 Elicitation of secondary metabolites in in
vitro cultures of Ammi majus L. Enz
19. Raskin, I., Ribnicky, D.M., Komarnytsky, S., Microbiol Technol. 33:565-568
Ilic, N., Poulev, A., Borisjuk, N., Brinker, A.,
Moreno, D.A. and Yakoby, R..N. (2002). 28. Xu, H., Kim, Y.K., Suh, S.Y., Udin, M.R.,
Plant and human health in the twenty-first Lee, S.Y. and Park, S.U. (2008). Deoursin
century. Trends Biotechnol. 20:522-531 production from hariy root culture of
Angelica gigas. J Korea Soc Appl Biol
20. Watson, A.A., Fleet, G.W.J., Asano, N., Chem. 51:349-351
Molyneux, R.J. and Nash, R.J. (2001).
Polyhydroxy latedalkaloid –natural 29. Kim, J.S., Lee, S.Y. and Park, S.U. (2008).
occurrence and therapeutic applications. Resveratol production in hairy root culture
Phytochemistry. 56:265-295 of peanut, Arachys hypogaea L. transformed
Saurabh Pagare et al
Current Trends in Biotechnology and Pharmacy 304
Vol. 9 (3) 293-304 July 2015, ISSN 0973-8916 (Print), 2230-7303 (Online)
with differet Agrobacterium rhizogenes 39. Zhao, J., Zhu, W. and Hu, Q. (2001).
strains. Afr J Biotechnol. 7:3788-3790 Enhanced catharanthine production in
Catharanthus roseus cell cultures by
30. Baldi, A. and Dixit, V.K. (2008). Enhanced
combined elicitor treatment in shake flasks
artemisinin production by cell cultures of
and bioreactors. Enz Microb Technol.
Artemisia annua. Curr. Terends Biotechnol
28:673-681
Pharmacol. 2:341-348
31. Olivira, A.J.B., Koike, L., Reis, F.A.M. and 40. Lee-Parsons, C.W.T. and Rogce, A.J.
Shepherd, S.L.K. (2001). Callus culture of (2006). Precursor limitations in methyl
Aspidosperma ramiflorum Muell.-Arg.: jasmonate-induced Catharanthus roseus
growth and alkaloid production. Acta cell cultures. Plant Cell Rep. 25:607-612
Scientia. 23:609-612
41. Ramani, S. and Jayabaskaran, C. (2008).
32. Sujanya, S., Poornasri, D.B. and Sai, I. Enhanced catharathine and vindoline
(2008). In vitro suspension cultures of production in suspension cultures of
Azadirachta indica. J Biosci. 33:113-120 Catharanthus roseus by ultraviolet-B light.
J Mol Signal. 3:9-14
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
cell suspension cultures of Brucea javanica suspension cultures of Cayratia trifoliata (L.)
(L.) Merr. Ind J Sci Technol. 1:1-6 Domin. Plant Biotechnol Rep. 3:135-138
34. Kusakari, K., Yokoyama, M. and Inomata,
43. Kim, O.T., Bang, K.H., Shin, Y.S., Lee, M.J.,
S. (2000) Enhanced production of
Jang, S.J., Hyun, D.Y., Kim, Y.C., Senong,
saikosaponins by root culture of Bupleurum
N.S., Cha, S.W. and Hwang, B. (2007).
falcatum L. using two step control of sugar
Enhanced production of asiaticoside from
concentration. Plant Cell Rep. 19:1115-1120
hairy root cultures of Centella asitica (L.)
35. Nikolaeva, T.N., Zagoskina, N.V. and Urban elicited by methyl jasmonate. Plant
Zaprometov, M.N. (2009). Production of Cell Rep. 26:1914- 1949
phenolic compounds in callus cultures of
tea plant under the effect of 2,4-D and NAA. 44. Kiong, A.L., Mahmood, M., Fodzillan, N.M.
Russ J Pl Physiol. 56:45-49 and Daud, S.K. (2005). Effects of precursor
supplementation on the production of
36. Umamaheswai, A. and Lalitha, V. (2007). triterpenes by Centella asiatica callus
In vitro effect of various growth hormones culture. Pak J BiolSci. 8:1160-1169
in Capsicum annum L. on the callus
induction and production of Capsiacin. J 45. Hohtola, A., Jalonen, J., Tolnen, A., Jaakola,
Plant Sci. 2:545-551 L., Kamarainen, T., Pakonen, M., Karppinen,
K., Laine, K., Neubauer, P., Myllykoshi, L.,
37. Nazif, N.M., Rady, M.R. and Seif, M.M.
Gyorgy, Z., Rautio, A. and Peltonen, O.
(2000). Stimulation of anthraquinone
(2005), Natural product formation by
production in suspension cultures of Cassia
plants, enhancement, analysis, processing
acutifolia by salt stress. Fitoterapia. 71:34-
and testing. In : Sustainable use renewable
40
natural resources – from principles to
38. Shrivastava, N., Patel, T. and Srivastava, practices (Eds. Jalkanen, A. and Nygren,
A. (2006). Biosynthetic potential of invitro P). University of Helsinki Publication. pp. 34-
grown callus cells of Cassia senna L. var. 69
senna. Curr Sci. 90:1472-1473
Secondary Metabolites of Plants and their Role
Current Trends in Biotechnology and Pharmacy 305
Vol. 9 (3) 293-304 July 2015, ISSN 0973-8916 (Print), 2230-7303 (Online)
46. Shohael, A.M., Chakrabarty, D., Yu, K.W., Gentianella austiaca shoot cultures. Plant
Hahn, E.J. and Paek, K.Y. (2005). Cell Tiss Org Cult. 94:329-335
Application of bioreactor system for large-
52. Mehrotra, S., Kukreja, A.K., Khanuja, S.P.S.
scale production of Eleutherococcus
and Mishra, B.N. (2008). Genetic
sessiliflorus somatic embryos in an air-lift
transformation studies and scale up of hairy
bioreactor and production of eleutherosides.
root culture of Glycyrrhiza glabra in
J. Biotechnol. 120: 228-236.
bioreactor. Elec J Biotechnol. 11:717-728
47. Schmeda-Hirschmann, G., Jordan, M.,
53. Gopi, C. and Vatsala, T.M. (2006). In vitro
Gertn, A., Wilken, D., Hormazabal, E. and
studies on effects of plant growth regulators
Tapia, A.A. (2004). Secondary metabolite
on callus and suspension culture biomass
content in Fabiana imbricate plants and in
yield from Gymnema sylvestre R.Br. Afr J
vitro cultures. Z Naturforsch. 5:48-54
Biotechnol. 5:1215-1219
48. Lee, S.Y., Cho, S.J., Park, M.H., Kim, Y.K.,
54. Misra, N., Misra, P., Datta, S.K. and
Choi, J.I. and Park. S.U. (2007). Growthand
Mehrotra, S. (2005). In vitro biosynthesis of
rutin production in hairy root culture of buck
antioxidants from Hemidesmus indicus
Downloaded From IP - 117.240.114.66 on dated 11-Aug-2016
Saurabh Pagare et al