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Triacontanol: A potent plant growth regulator in agriculture

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Triacontanol: a potent plant growth regulator in

agriculture
a a a
M. Naeem , M. Masroor A. Khan & Moinuddin
a
Department of Botany, Plant Physiology Section, Aligarh Muslim University, Aligarh, 202
002, Uttar Pradesh, India

Available online: 06 Sep 2011

To cite this article: M. Naeem, M. Masroor A. Khan & Moinuddin (2012): Triacontanol: a potent plant growth regulator in
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 Journal of Plant Interactions
Vol. 7, No. 2, June 2012, 129
142

REVIEW ARTICLE
Triacontanol: a potent plant growth regulator in agriculture
M. Naeem*, M. Masroor A. Khan and Moinuddin

Department of Botany, Plant Physiology Section, Aligarh Muslim University, Aligarh 202 002, Uttar Pradesh, India
(Received 16 May 2011; final version received 29 August 2011)

Triacontanol (TRIA) is a natural plant growth regulator found in epicuticular waxes. It is used to enhance the
crop production in millions of hectares, particularly in Asia. Quite a number of researchers have reported the
TRIA-mediated improvement in growth, yield, photosynthesis, protein synthesis, uptake of water and nutrients,
nitrogen-fixation, enzymes activities and contents of free amino acids, reducing sugars, soluble protein, and active
constituents of essential oil in various crops. Expectedly, TRIA enhances the physiological efficiency of the cells
and, thus, exploits the genetic potential of plant to a large extent. In fact, TRIA increased free amino acids,
Downloaded by [T&F Internal Users], [Rob Blackmore] at 02:45 17 May 2012

reducing sugars, and soluble protein of rice (Oryza sativa L.) and maize (Zea mays L.) within 5 min. TRIA elicited
the appearance of L()-adenosine within 1 min in the roots of plants, the shoots of which were sprayed with
nanomolar concentrations of TRIA. TRIA and octacosanol (OCTA), the primary alcohols, are ubiquitous in the
environment. OCTA was reported to inhibit the activity of TRIA in the seedlings of rice (Oryza sativa L.) at
equimolar concentrations; and both TRIA and OCTA elicited a second messenger, known as OCTAM and
triacontanol second messenger (TRIM), respectively. TRIA rapidly increases the ratio of L()- to D(
)-
adenosine, probably at the tonoplast. However, it is to be resolved as to how TRIA elicits L()-adenosine and
what is the source of L()-adenosine in plants. Based on known metabolic processes, de novo synthesis of L()-
adenosine is unlikely, because of the rapidity of the response. TRIA-mediated increase in dry matter production
could influence the inter-relationship between primary and secondary metabolism, leading to increased
biosynthesis of secondary products. Various studies present strong evidences that application of TRIA applied
either to the root medium or to leaves enhanced the growth and yield of vegetables and other crops, including
agronomic and horticultural crops as well as medicinal and aromatic crop plants under normal and adverse
conditions. However, further investigations are required to elucidate the possible role of TRIA on plant growth
regulation, physiological activities and secondary metabolite biosynthesis regarding medicinal and aromatic
plants subjected to abiotic stress. The present review covers the pivotal role of TRIA in plant growth and
development, its mode of action and its significance in improving the crop productivity and quality of agricultural
crops.

Keywords: medicinal and aromatic crops; plant growth; crop yield; bioactive constituents; triacontanol

Introduction Ogasawara 1981). The callus tissue contained a

The plant growth regulatory activity of triacontanol homologous series of alkanes, namely. n-C23H48,
(TRIA) was first discovered by Ries et al. (1977) in n-C25H52, and n-C27H56 compared to n-C27H56,
alfalfa (Medicago sativa L.). According to Mandava n-C29H60, and n-C31H64 found in the rice leaves.
(1979), TRIA is a secondary plant growth substance This observation might explain as to why TRIA could
and cannot be considered as a phytohormone. Such enhance callus growth in cultures of different species
types of growth regulators enhance the physiological (Hangarter et al. 1978).
efficiency of the cells and, thus, exploit the plant Many researchers have reported the positive role
genetic potential to a large extent. Earlier studies have of TRIA in enhancing growth, yield, photosynthesis,
given sound evidences regarding distribution of nitrogen fixation, enzymes activities, free amino
TRIA in the epicuticular waxes in widely diverse acids, reducing sugars, and soluble protein of plants
genera, such as California Croton (Croton (Ries 1991; Ries et al. 1993; Nagoshi and Kawashima
californicus), blueberry (Vaccinium ashei), Brazilian 1996; Borowski et al. 2000; Naeem et al. 2009; Aftab
palm (Copernica cerifera), runner bean (Phaseolus et al. 2010; Idrees et al. 2010; Naeem et al. 2010,
multiflorus), white clover (Trifolium repens), alfalfa Naeem et al. 2011). Muthuchelian et al. (1997, 2003)
(Medicago sativa), and in physic nut (Jatropha curcas) investigated the significant effect of TRIA on acid
(Hufford and Oguntimein 1978; Luzbetak et al. 1978; mist-treated Erythrina variegata L. seedlings with
Freeman et al. 1979; Lee et al. 1979). TRIA is particular emphasis on growth, photosynthetic pig-
considered to be the only primary alcohol (Figure 1) ments, ribulose-1,5-bisphosphate (RuBPC), and
found in the wax of the rice leaves (Uchiyama and photosynthetic activity. Kumaravelu et al. (2000)

*Corresponding author. Email: [email protected]; [email protected]

ISSN 1742-9145 print/ISSN 1742-9153 online
# 2012 Taylor & Francis
http://dx.doi.org/10.1080/17429145.2011.619281
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 130 M. Naeem et al.

OH labs), were active at extremely low concentrations (23

nM). TRIA also promoted growth of tobacco callus
at concentrations as low as 0.01 mg per Petri dish
when applied to either roots or shoots (Ries and Wert
1977). As suggested by Jones et al. (1979) and Ries
and Houtz (1983), there are possibilities that impu-
Triacontanol (TRIA) rities in TRIA might inhibit the physiological effect of
TRIA at higher levels. Application of even pico mole
Figure 1. Structural formula of triacontanol (TRIA).
quantities of ()-adenosine (a precursor in TRIA
biosynthesis and a plant growth-promoting second
reported the ameliorating effect of TRIA applied at
0.5 mg dm
3 on growth and yield attributes, photo- messenger) to the foliage of tomato (Lycopersicon
synthetic pigments, nitrate reductase (NR) activity esculentum Mill.), maize (Zea mays L.), and cucum-
and contents of nitrate, protein, starch, amino acids, ber (Cucumis sativa L.) increased the concentrations
and total phenols in green gram (Vigna radiata L.). of Ca2  , Mg2  , and K by 20
60% in the exudates
TRIA application increased the plant dry weight, obtained from the stumps of excised plants within 5
protein, and chlorophyll contents and net photosyn- seconds of TRIA application (Ries et al. 1993).
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thetic rate in rice (Oryza sativa L.) (Chen

et al. 2002). Similarly, exogenous application of
TRIA showed positive effect on growth, chlorophyll Mode of TRIA action
content, efficiency of photosystem-II (PS-II) and gas Many investigators have explored the effects of TRIA
exchange characteristics in the seedlings of rice on several basic metabolic processes including photo-
(Muthuchelian et al. 1995; Kumaravelu et al. 2000), synthesis, nutrient uptake, and enzyme activities.
maize (Ries 1991), and wheat (Ries 1991). Increase in Several efforts have been made to elucidate the
plant growth could mainly be due to an abrupt mechanism of TRIA action (Ries and Houtz 1983;
TRIA-induced increase in photosynthesis as TRIA Ries 1991; Ries et al. 1993). Assumption of a cascade
has been reported to be involved in the up-regulation effect led to the identification of 9-b-L ()-adenosine
of many genes involved in the photosynthetic process as a second messenger of TRIA (Ries 1991; Ries et al.
(Chen et al. 2002, 2003). 1993). TRIA rapidly elicits the second messenger
The present review emphasizes the role of TRIA (TRIM) in rice (Oryza sativa L.), which at nanomolar
in enhancing growth, yield, and quality of crop concentrations causes the plants to respond in a
plants as well as in improving physiological and
manner similar to TRIA (Ries 1991). TRIM has
biochemical attributes and active constituents of
been identified as 9-b-l ()-adenosine (9H-purin-6-
medicinal and aromatic plants under normal and
amine, 9-b-L-ribofuranosyl) as depicted in Figure 2.
stressful conditions.
TRIA enhanced the formation or release of L()-
adenosine in the root tissue of rice seedlings within one
Physiological activity of TRIA at low concentrations min of TRIA application to the shoots, which might
have elucidated the first step in the mode of TRIA
According to Ries and Houtz (1983), the low action (Ries et al. 1990). It is already reported that
concentrations of TRIA might be biologically active TRIA rapidly increases the ratio of l()-adenosine to
because of the sensitivity of the plants to its extremely d(
)-adenosine, probably at the tonoplast (Ries 1991).
low doses. The physiological activity of TRIA in There remains the problem of how TRIA elicits l()-
maize (Zea mays) and rice (Oryza sativa) seedlings adenosine and what is the source of l()-adenosine in
has been improved consistently by increasing the plants. Based on known metabolic processes, de novo
purity of TRIA to 99.7% (Ries and Wert 1982). As
synthesis of l()-adenosine is unlikely, because of the
per earlier studies of Ries and Wert (1977), TRIA is
rapidity of the response. The most probable source of
active at concentrations as low as 10 mg L
1 (2.310-8
adenosine is AMP derived from ADP and ATP
M) when applied either to shoots or roots of a crop.
(Olsson and Pearson 1990). In TRIA treated plants,
Since TRIA is extremely insoluble in water (2 10-16
M or 910
14 g L
1), the solution of TRIA has been
formulated as a stable colloidal dispersion of fine
crystalline particles in water (Laughlin et al. 1983).
This formulation indicates accurate quantification of
the application rate and a smooth dose-response
curve for dry weight gain of corn as a result of
TRIA application. The TRIA, applied at 10 mg L
1,
increased the dry weight and leaf area of rice plants in
nutrient cultures (Ries and Wert 1977). Similarly,
Hangarter et al. (1978) found that TRIA, isolated
from alfalfa or the one synthesized artificially (Ana- Figure 2. Structural formula of adenosine.
 Journal of Plant Interactions 131

the nonracemic L(
)-adenosine (
11%) is released to soluble proteins, reducing sugars, and free amino
affect plant processes (Ries 1991). acids (Ries 1985, 1991). Similar increases in these
Ries et al. (1993) in their study, noticed that more metabolic constituents occurred in the cell-free ex-
information is needed regarding protocols that will tracts of maize leaves treated with TRIA, although
ensure consistent results with both TRIA and L()- the response was less rapid (Ries 1985). The rapid
adenosine in growth chamber, greenhouse, and field increase in total nitrogen by TRIA treatment is also
studies. However, the strong evidence is overwhelm- difficult to explain. This increase, whether expressed
ing that TRIA applied at nanomolar concentrations in total N or as soluble N and insoluble N, was
might increase the yield or improve the quality of magnified when there was an accompanying increase
various plant species. TRIA increased the dry weight in dry weight of TRIA treated plants (Knowles and
as well as content of free amino acids, reducing Ries 1981). A hypothesis is that plant metabolism is
sugars, and soluble protein of rice (Oryza sativa L.) altered by TRIA or its second messenger L()-
and maize (Zea mays L.) plants within 5 min (Ries adenosine, leading to expected compositional or
1991). TRIA also elicited the appearance of L()- chemical changes (Diehi 1974; Knowles and Ries
adenosine in the roots of plants whose shoots were 1981). The elicitation of the second messenger of
sprayed with its nanomolar concentrations within 1 TRIA (L()-adenosine) in the roots within 1 min
min (Ries and Wert 1988). This was the first evidence after TRIA application to the foliage is also difficult
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that L()-adenosine occurred in nature. Synthetic to explain based on our current knowledge of
L()-adenosine increased the rate of growth of rice translocation of substances and enzyme kinetics.
seedlings, as measured by total dry weight gain of The L()-adenosine had the same effect on plants
more than 50% within 24 h at foliar application rate as TRIA, and apparently it did not elicit any other
of 0.01
100.0 mg L
1 (3.7 10
11 to 10
7 M); whereas, chemical compound which stimulated the plant
D(
)-adenosine did not affect the plant growth at all growth (Ries et al. 1990, 1993).
(Ries 1991). In relation to the mode of action of
exogenously applied TRIA and L()-adenosine, it is
postulated that both of the compounds move rapidly Response of plants to TRIA
through the leaf cuticle to the plasma membrane of
Growth attributes
epidermal cells of tomato (Lycopersicon esculentum
Mill.), maize (Zea mays L.), and cucumber (Cucumis Various studies provided strong evidence that appli-
sativa L.). TRIA then elicits the formation of l()- cation of TRIA, applied either through the root
adenosine. The study suggested that L()-adenosine medium or to the leaves, enhanced the growth and
triggered a rapidly transmitted signal within whole yield of vegetables and cereal crops (Ries et al. 1977,
plants that resulted in a transient increase in apo- 1978). TRIA has shown to increase the growth and/or
plastic ion concentration within stem tissue (Ries et al. yield of most of the major annual vegetables,
1993). agronomic and horticultural crops as well as forest
Thus, it is also a significant challenge to discover species (Ries 1985, 1991; Kawashima et al. 1987;
as to how both TRIA and L()-adenosine enhance Kapitsimadi and Vioryl 1995). The increase in yield is
plant metabolism and how TRIA elicits L()-ade- due to the rapid increase in the net assimilation rate
nosine. Further, identification of octacosanol second as observed in tomato after TRIA spray (Ries 1985,
messenger (OCTAM) and the determination of how it 1991). In greenhouse studies, foliar applications of
inhibits the activity of TRIA may lead to discovery of 1.0
100 mg L
1 of L()-adenosine increased the
the site of TRIA and/or L()-adenosine action (Ries growth of tomato (Lycopersicon esculentum Mill.),
1991; Ries et al. 1993). maize (Zea mays L.), cucumber (Cucumis sativus L.),
and carrot (Daucus carota L.) (Ries et al. 1990).
Further, in controlled environment studies, Eriksen
TRIA elicits L()-adenosine et al. (1982) noticed a TRIA-mediated increase in the
Adenosine (abbreviated as TRIM) was found to be a dry weight of tomato (Lycopersicon esculentum)
naturally occurring plant growth substance elicited by seedlings; but, such a dry weight increase was not
TRIA (Ries et al. 1990). The adenosine from TRIA- observed in maize seedlings. TRIA showed no
treated plants was identical to the adenosine obtained effect on seed germination or on early growth of
from control plants as determined by melting point, several species when seeds were treated with it
IR and MS, and from proton and carbon NMR (Hoagland 1980), but significant effect of TRIA was
analysis (Ries et al. 1990). However, only the pure reported in increasing the rate of germination of
adenosine obtained from TRIA-treated plants could cotton (Gossypium hirsutum) (Zerong et al. 1981) and
stimulate the plant growth. Other forms of adenosine, leguminous crops (Janardhan 1992). Skogen et al.
including d(
)-adenosine, did not stimulate growth at (1982) reported increases in growth, number
similar concentrations. Thus, it was postulated that of inflorescences, and quality of flowers treated
the adenosine from TRIA-treated plants was L()- with TRIA in chrysanthemum (Chrysanthemum
adenosine, the enantiomer of d(
)-adenosine. In morifolium). Foliar spray of TRIA increased the dry
addition, TRIA caused simultaneous increases in weight of rice seedlings grown in nutrient medium
 132 M. Naeem et al.

and that of the plants of corn, barley, and tomato Triacontanol increased the minimal fluorescence
(Ries et al. 1977). The growth promoting effects of and the maximal fluorescence (Fo and Fm) in rice
TRIA on various attributes, especially on plant plants, as revealed by Chen et al. (2003). They
height, fresh and dry weights, leaf-area and root explained that increased Fo value indicated that
nodulation, have been explored in various medicinal TRIA increased the antenna pigment level or the
crops by various researchers (Srivastava and Sharma efficiency of excitation trapping at the active centers
1990; Misra and Srivastava 1991; Shukla et al. 1992; of PS-II. These data were similar with those obtained
Muthuchelian et al. 2003; Giridhar et al. 2005; in tomato (Borowski et al. 2000). They further
Chaudhary et al. 2006). TRIA improved fresh and reported a quantitative increase (30% increase over
dry weights, leaf area, and the number and dry weight the control) in RuBisCO as a result of TRIA
of nodules in case of hyacinth bean (Lablab purpureus application. Other proteins were unaffected quantita-
L.) (Naeem and Khan 2005; Naeem et al. 2010). tively by TRIA treatment.
Furthermore, the effect of foliar spray of GA3, TRIA, Photosynthesis has been implicated as an impor-
and that of their combination (GA3TRIA) was tant plant response to TRIA (Eriksen et al. 1981) and
found to be significant for plant height and dry the increased growth and dry weight of plants were
weight of opium poppy (Khan et al. 2007). Applica- attributed to improved photosynthesis and enhanced
tion of TRIA alone and in combination with potas- accumulation of photosynthates. A number of studies
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sium enhanced the plant height, fresh and dry weights have demonstrated an increased rate of CO2 fixation
and area per leaf of tomato (Khan et al. 2009). in a variety of plant species when the TRIA was
Various studies, conducted on the medicinal and applied in nanomolar concentrations (Haugstad et al.
aromatic plants (Table 1), revealed that foliar sprays 1983; Popova et al. 1989; Srivastava and Sharma
of TRIA significantly stimulated the growth attri- 1990; Misra and Srivastava 1991). Houtz et al.
butes of turmeric (Curcuma longa L.), hyacinth bean (1985b) suggested a TRIA-stimulated increase in the
(Lablab purpureus L.), ashwagandha (Withania specific activity of the enzymes RuBisCO and phos-
somnifera L.), artemisia (Artemisia annua L.), cor- phoenolpyruvate carboxylase. Moreover, TRIA af-
iander (Coriandrum sativum L.), coffee senna (Senna
fected several other enzyme systems especially
occidentalis L.), mint (Mentha arvensis L.), and ginger
the membrane bound Ca2  /Mg2  dependent AT-
(Zingiber officinale Rosc.) (Singh 2008; Naeem
Pase (Lesniak et al. 1986, 1989). TRIA also increased
et al. 2009; Nasir 2009; Aftab et al. 2010; Idrees
the activity of the key respiratory enzyme malate
et al. 2010; Naeem et al. 2010, Naeem et al. 2011;
dehydrogenase (Savithiry et al. 1992). Ries and Houtz
Singh et al. 2011).
(1983) suggested that the hormonal effects of TRIA
might be due to TRIA-induced alterations at the
Physiological and biochemical attributes membrane level. TRIA-mediated activation of a
number of membrane bound enzymes supports this
Exogenous application of TRIA has been reported to
regulate directly or indirectly several physiological assumption (Ries and Wert 1988; Ries 1991; Savithiry
and biochemical processes (Ries and Houtz 1983; et al. 1992). As per Ries et al. (1990), TRIA treatment
Ries 1991; Ries et al. 1993; Naeem et al. 2009, 2010, causes rapid elicitation of a specific second messen-
2011). Application of TRIA to barley roots resulted ger, 9-b-L() adenosine, which could induce extre-
in a rapid stimulation of membrane associated Ca2  / mely rapid physiological responses.
Mg2  dependent ATPase activity (Lesniak et al. Ivanov and Angelov (1997) observed a significant
1986) in a calmodulin-dependent manner. Besides, increase of the net CO2 uptake to 109% and 119%,
activity of NADPH oxidase of the plasma membrane when the TRIA was applied to pea plants at 10
8 and
is potentiated by TRIA application (Moore et al. 10
6 M, respectively. Besides, they noted that treat-
1991). Isolation and characterization of TRIA-regu- ments with octacosanol (OCTA) under the same
lated genes was the first step toward understanding experimental conditions and concentrations did not
the TRIA action, since it gave clues to the biochem- elicit any changes in photosynthesis over the same
ical pathways and physiological processes that reg- time range (30 and 60 min). Ries and Houtz (1983)
ulate, and reveal the components involved in TRIA suggested for the first time that the growth stimulating
signaling (Chen et al. 2002). A large number of the or hormonal effects of TRIA might be due to certain
TRIA responsive genes were associated with photo- alterations at the cell membrane level. They demon-
synthesis. These genes were upregulated and the stress strated that net photosynthesis was stimulated to a
related genes were down regulated by TRIA. Chen et greater extent in isolated protoplast preparations,
al. (2002) reported that higher rbcS gene levels were supporting the suggestion that the putative initial
associated with improved photosynthetic activity in site of TRIA action could be localized at the level of
TRIA-treated plants. They also illustrated that plasma membranes. As per their study, Ivanov and
TRIA affected the photosynthesis by increasing the Angelo (1997) suggested that TRIA-induced increase
level and activity of ribulose-1,5-bisphosphate in the content of pyrene excimer (excitation light beam
carboxylase oxygenase (RuBisCO) and by improving [steady state fluorescence spectra]) was due to the
the status of photosystems (Chen et al. 2003). molecular dynamics and/or fluidity of protoplast and
 Table 1 Positive response of various plant species to foliar application of triacontanol.

Name of plant Botanical name Family name Growth attributes Yield attributes Biochemical attributes Quality attributes Reference citation

Opium poppy Papaver Papaveraceae Plant height, dry Number of capsules, Chl a, Chl b, and total content Morphine content and Khan et al. (2007)
somniferum L. weight and number seed yield per plant, morphine yield per plant
of branches and crude opium
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yield per plant

Tomato Solanum Solanaceae Height per plant, Number of fruits per Total chl and carotenoids Fruit ascorbic acid and Khan et al. (2009)
lycopersicum L. number of leaves and plant, weight per fruit content, leaf-N, -P, and -K lycopene contents
plant fresh and dry and fruit yield per contents
weights plant
Hyacinth bean Lablab purpureus Fabaceae Plant fresh and dry Number of pods per Photosynthetic rate (PN), Seed-protein content, Naeem and Khan
L. weights, leaf-area per plant, number of seeds stomatal conductance (gs) total carbohydrate (2005), Naeem et
plant, number and dry per pod, 100-seed and transpiration rate, total content, and tyrosinase al. (2009)
weights of nodules weight and seed-yield chl and carotenoid content, activity

Journal of Plant Interactions

per plant NR and CA activities, leaf-N, -
P, -K, and -Ca content,
nodule-N and leghemoglobin
contents
Artemisia Artemisia annua Asteraceae Shoot and root lengths, Artemisinin yield PN, gs and transpiration rate, Essential oil content, Aftab et al. (2010)
L. plant fresh and dry total chl and carotenoid artemisinin content
weights content, NR and CA activities,
leaf-N, -P, and -K content
Coriander Coriandrum Umbelliferae Shoot and root lengths, Total chl and carotenoids Essential oil content Idrees et al. (2010)
sativum L. plant fresh and dry content, NR and CA activities,
weights leaf-N, -P, and -K content
Coffee senna Senna Fabaceae Plant fresh and dry Number of pods per PN, gs and transpiration rate, Total anthraquinone Naeem et al.
occidentalis L. weights plant, number of seeds total chl and carotenoid and sennoside contents, (2010)
per pod, 100-seed content, NR and CA and seed-protein content
weight and seed yield activities, leaf-N, -P, -K,
per plant and -Ca content
Sweet basil Ocimum Labiatae Shoot and root lengths, Essential oil yield Chl a, Chl b, total Chl, and Leaf-protein and Hashmi et al.
basilicum L. number of spikes per carotenoid contents, activities carbohydrate contents, (2011)
plant, total leaf area, of NR and CA, leaf-N, -P, essential oil content,
plant fresh and dry and -K contents linalool, methyl eugenol,
weights and eugenol contents
Japanese mint Mentha arvensis Lamiaceae Plant height, leaf-area, Herbage yield, Total chl and carotenoid Essential oil content, Naeem et al.
L. leaf-yield, and plant essential oil yield contents, activities of NR menthol, L-menthone, (2011)
fresh and dry weights and CA, leaf-N, -P, and -K isomenthone, and
contents, total phenol menthyl aceteate
contents

133
Abbreviation: Chl, Chlorophyll; NR, Nitrate reductase; CA, Carbonic anhydrase.
 134 M. Naeem et al.

chloroplast membranes that were markedly enhanced poppy (Khan et al. 2007), hyacinth bean (Naeem
as a result of TRIA application. Furthermore, they et al. 2009), artemisia (Aftab et al. 2010), coriander
noted that the concentration range of TRIA, effective (Idrees et al. 2010), and coffee senna and mint
in the enhancement of photosynthesis, corresponded (Naeem et al. 2010, 2011).
well to the TRIA concentrations influencing mem- Triacontanol application caused an increase in
brane dynamic properties; this indicated a good peroxidase activity in ‘Little Marvel dwarf’ (LM) and
correlation between the two phenomena. They also ‘Alaska’ peas (AP) plants compared to the untreated
demonstrated that the incubation of both protoplast controls. The effects of TRIA on root and stem
and chloroplasts with TRIA resulted in a rise of the growth, peroxidase activity, and auxin-destruction
excimer/monomer (IE/IM) ratio of pyrene fluores- appeared to be cultivar-specific, with respect to LM
cence. These results are discussed in terms of specific and AP varieties of peas (Henry and Gordon 1980).
concentration-dependent TRIA-induced alterations As a result of foliar spray of TRIA at early vegetative
of the dynamic properties of protoplast and chlor- stages, Naeem et al. (2009) reported an increase in the
oplast membranes and their possible involvement in nodule-nitrogen and leghemoglobin contents of hya-
the initiator and integral physiological response to cinth bean (Lablab purpureus L.) at 60 days after
exogenous application of TRIA (Ivanov and Angelov sowing (DAS) followed by that at 90 DAS. The
1997). In a study, Borowski et al. (2000) revealed that TRIA applied at 10
6 concentration surpassed the
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the maximum efficiency of PS-II in the dark (Fv/Fm) control by 5.4% in nodule-nitrogen content and by
was clearly increased by the applied TRIA, and the 14.1% in leghemoglobin content at 60 DAS. How-
efficiency of excitation-capture by PS-II reaction ever, the plants gave negative response to 10
5 M of
centers (Fv/Fm) were even much increased. TRIA both for nodule-nitrogen and leghemoglobin
In fact, a number of studies have demonstrated an contents in comparison to 10
6 M of TRIA.
increased rate of CO2 fixation and photosynthesis in a The significant effect of TRIA on activities of NR
variety of plant species as a result of TRIA applica- and carbonic anhydrase (CA) has been reported by
tion in nanomolar concentrations (Houtz et al. 1985a; Kumaravelu et al. (2000), Muthuchelian et al. (2003),
Ivanov and Angelov 1997). Srivastava and Sharma and Nasir (2009). The TRIA alone or in combination
(1990) and Misra and Srivastava (1991) reported with gibberellic acid significantly enhanced the activ-
significant improvement in the rate of CO2 fixation ities of NR and CA in artemisia (Aftab et al. 2010)
and net photosynthesis in opium poppy (Papaver and coriander (Idrees et al. 2010). Moreover, a
somniferum L.) and lemongrass (Cymbopogon remarkable enhancement in the activities of NR and
flexuosus Steud. Wats.), respectively. According to CA in hyacinth bean, coffee senna and mint was
Muthuchelian et al. (2003), when the seedlings of reported by Naeem et al. (2009, 2010, 2011) as a result
Erythrina variegata L. were sprayed with TRIA, the of TRIA application.
harmful effect of acidic mist on photosynthesis Naeem et al. (2009) reported significant effect of
machinery was partially or completely reversed, foliar application of TRIA on N, P, K, and Ca
indicating that TRIA might protect plants from content of leaves in hyacinth bean. The concentra-
acid rain. TRIA application also improved the tions of these leaf-nutrients were found significantly
photosynthesis in rice in normal conditions (Chen et higher at 10
6 M of TRIA over the control. There-
al. 2002). Similarly, TRIA alone or in combination after, at 10
5 M of TRIA, the tissue concentrations of
with gibberellic acid (1.5 mg L
1 of TRIA75 mg L
1 these elements decreased. A positive effect of TRIA
of GA3) significantly increased the net photosyn- on total leaf-nitrogen content was earlier revealed by
thetic rate, stomatal conductance and internal CO2 Knowles and Ries (1981) and Ries and Houtz (1983)
concentration in Artemisia annua (Aftab et al. 2010). in the case of rice and maize, respectively. Knowles
Moreover, Naeem et al. (2009, 2010, 2011) also found and Ries (1981) clarified that enhancement in leaf-
the significant effect of TRIA on photosynthetic nutrients, particularly that in nitrogen, due to TRIA
parameters regarding hyacinth bean (Lablab application could be attributed to the compositional
purpureus L.), coffee senna (Senna occidentalis L.), or chemical change in plants leading to alterations in
and Japanese mint (Mentha arvensis L). leaf-nitrogen concentration. In contrast, TRIA ap-
In addition, the contents of photosynthetic pig- plied at 0.5 and 1.0 mg dm
3, reduced the leaf nitrate
ments were significantly influenced by exogenous content in green gram (Kumaravelu et al. 2000). A
application of TRIA. The content of the pigments significant improvement in leaf-nutrient content of
in TRIA-treated leaves could presumably be attrib- tomato, opium poppy, artemisia, coriander, and
uted to the increase in the number and size of ginger was observed by Khan et al. (2006, 2007,
chloroplasts as revealed by Ivanov and Angelov 2009), Aftab et al. (2010), Idrees et al. (2010), and
(1997), Chen et al. (2003), and Muthuchelian et al. Singh et al. (2011) as a result of TRIA application
(2003). Significant effects of TRIA on photosynthetic alone or in combination with GA3. TRIA application
pigments were also reported in opium poppy and at 10
6 M alleviated the leaf-N, -P, and -K contents in
Japanese mint (Srivastava and Sharma 1990, 1991), the case of coffee senna and mint, respectively
lemongrass (Misra and Srivastava 1991), opium (Naeem et al. 2010, 2011). Sharma et al. (2002)
 Journal of Plant Interactions 135

suggested that a higher content of leaf-nutrients in Kawashima et al. 1987). Borowski et al. (2000)
TRIA treated plants could be attributed to the higher reported significant increase in tomato yield as result
metabolic activity and increased dry matter produc- of TRIA application at 0.3 and 3.0 mg L
1. Similar
tion that might result in enhanced water and nutrient results have been reported by Asane et al. (1998),
uptake from soil subsequently. Barus (1998), and Blamowski et al. (1998) on peas,
plum, and radish, respectively. In another study,
Eriksen et al. (1982) noticed a significant increase in
Yield and quality attributes total as well as per plant yield of tomato, when TRIA
Past decades have witnessed much success in increas- was applied as foliar sprays; but when TRIA was
ing the yield of food crops and vegetables using TRIA added to the growth medium, only a temporary
(Bouwkamp and McArdle 1980; Eriksen et al. 1981; increase in yield and number of fruits was observed.
Ries and Houtz 1983; Hashim and Lundergan 1985; Further, they found no effect of TRIA on maize yield,
Ries 1985; Stoutemyer and Cooke 1987; Borowski whether the TRIA was applied to the leaves or it was
et al. 2000; Muralidharan et al. 2000). The first added to the growth substrate. Verma et al. (2009)
published report on TRIA showed a significant maintained that TRIA did not induce enhancement in
increase in yield on several crops including dry beans, pod yield, number of pods per plant, pod weight per
sweet corn, and cucumbers (Ries et al. 1978; Ries and plant, and shelling percentage of groundnut. There
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Houtz 1983). TRIA, applied through soil drenching, were recorded the highest values regarding the
has been shown to be effective in promoting the number of umbels, fruits per umbel, 100 seed weight,
growth and development of tabasco pepper both in and seed yield, when the TRIA was applied together
greenhouse and field conditions (Mamat et al. 1983). with GA3 to coriander crop (Idrees et al. 2010)
Bhalla (1981) reported the TRIA-enhanced growth of Naeem et al. (2009, 2010) reported a favorable
maize and tomato plants grown in the greenhouse. effect of TRIA on the yield and yield attributes
The results of 46 field experiments, conducted in (number of pods and seed yield per plant) of hyacinth
several parts of the world, generally showed no
bean and coffee senna; the highest values were
significant increases in crop yield, except one study
recorded when the TRIA was applied at a 10
6 M
conducted in Japan, where the yield of rice was
concentration. Ivanov and Angelov (1997), Borowski
increased by 17
21% on account of soil applications
et al. (2000), Kumaravelu et al. (2000), Chaudhary
of TRIA at 0.057
4.0 g ha
1. However, foliar and
et al. (2006), Sharma et al. (2006), and Nogalska et al.
seed treatment with TRIA at 0.05 and 0.10 mg L
1
(2008) reported the significant effect of TRIA on the
increased the yield of cotton by 12 and 31%,
yield and yield attributes of pea, tomato, green gram,
respectively (Sheng 1981). The average yield of winter
water chestnut, tomato, and soybean, respectively. A
wheat was increased by 12% due to foliar application
significant improvement in fruit-yield of tomato
of TRIA at 0.1 and 0.5 mg L
1 (Chen et al. 1980). As
(Khan et al. 2006, 2009) and crude opium-yield, and
per an extensive study in China conducted in a three-
year period, several vegetable crops (tomato, egg- seed yield of opium (Khan et al. 2007) has also been
plant, cabbage, and winged bean) responded posi- reported as a result of TRIA application carried out
tively to 0.1
1.0 mg L
1 of TRIA. However, in the alone or in combination with GA3. Foliar application
USA, TRIA increased the plant growth in labora- of TRIA at 10
6 M significantly enhanced the crop
tory-study as well as in greenhouse-study; however, herbage yield and the yield of essential oil in mint
results obtained in the field study were not encoura- (Naeem et al. 2011). Application of TRIA (at 10
6 M)
ging (Ries and Houtz 1983). also improved the values of primary and secondary
In brown rice, Nagoshi and Kawashima (1996) fingers and rhizome yield per plant in turmeric and
reported a TRIA-mediated improvement in heading ginger (Singh 2008; Singh et al. 2011). According to
and percentage of ripened grain at harvesting time Nasir (2009), TRIA combined with N and P fertilizers
that led to the enhancement in the individual grain (10
6.5 M of TRIA N60P40) significantly increased
weight and 1000-grian weight. Chowdhury et al. the capsules per plant and seed yield of datura
(2009) found the beneficial effect of TRIA in improv- (Datura innoxia Mill.). The application of TRIA in
ing fruit quality and economic yield of different combination with N and P (10
6.5 M of
cultivars of water chestnut (Trapa bispinosa); TRIA TRIA N45P26) also increased the seed yield and
application increased the fresh fruit yield by 32% in root yield of withania (Withania somnifera L.) (Nasir
green cultivars and by 31% in red cultivars. Foliar 2009).
spray of 0.5 mg dm
3 of TRIA significantly promoted Foliar application of TRIA, at a concentration of
the onset of flowering and increased the pod produc- 0.5 mg dm
3, significantly promoted the contents of
tion pod number, seed number, mass per plant, and saccharides, starch, soluble proteins, amino acids,
mass per pod in green gram plants exposed to TRIA and phenols in green gram (Kumaravelu et al. 2000).
at 0.5 mg dm
3 (Kumaravelu et al. 2000). Improve- TRIA application also improved the contents of
ment in yields of several important food crops have soluble protein, starch, sugars, and free amino acids
been recorded by several researchers as a result of in the leaves of Oryza sativa and Zea mays (Kim et al.
TRIA application (Ries et al. 1978; Ries 1985; 1989), Acacia catechu (Thakur and Thakur 1992), and
 136 M. Naeem et al.

Erythrina variegata (Muthuchelian et al. 1995). More- significantly improved the accumulation of essential
over, Naeem et al. (2009, 2010) reported a significant oil and its components, particularly that of citral-a
positive effect of TRIA on the seed-content of protein and citral-b; whereas, TRIA had no effect on the
and carbohydrate in hyacinth bean. Similarly, carbo- geraniol content of the oil. As a result of foliage-
hydrate and protein contents in turmeric (Curcuma applied TRIA, Farooqi and Sharma (1988) found an
longa L.) and ginger (Zingiber officinale Rosc.) were increase in the essential oil yield in Mentha arvensis.
significantly improved by the foliar spray of TRIA Srivastava and Sharma (1991) reported an improved
(Singh 2008; Singh et al. 2011). content of menthol and yield of essential oil in TRIA-
treated M. arvensis plants. Aftab et al. (2010) noted a
significant enhancement in the content of Artemisia
Active constituents of plants
annua, registering a simultaneous increase in
Khan et al. (2007) found a significant positive effect the content and yield of artemisinin in the leaves
of cumulative application of TRIA and GA3 on the (Figure 3). Similarly, Idrees et al. (2010) revealed the
yield of opium and its morphine content. The crude- significant positive effect of combined application of
opium production of opium poppy (Papaver TRIA and GA3 on the essential oil content of
somniferum L.) was also enhanced due to the com- coriander (Figure 4). Moreover, Naeem et al. (2010)
bined application of TRIA and GA3 (Khan et al. recorded a TRIA-mediated improvement in the level
Downloaded by [T&F Internal Users], [Rob Blackmore] at 02:45 17 May 2012

2007). Srivastava and Sharma (1990) also examined of anthraquinone and sennoside content in coffee
the effect of TRIA on alkaloid-biosynthesis as well as senna (Figure 5). In addition, Naeem et al. (2011)
on the relationship between alkaloid production and recorded a TRIA-mediated improvement in the
physiological parameters in opium poppy. They contents of components of essential oil (menthol, l-
reported a significant increase in capsule number methone, isomenthone, and menthyl acetate) of
and morphine content of the plant owing to foliar
Mentha arvensis L. (Figure 6). Total alkaloid content
application of TRIA at concentration of 0.01 mg L
1;
in withania (Withania somnifera L.) and datura
whereas, there was no effect of TRIA on thebaine and
(Datura innoxia Mill.) and curcumin content of
codine contents of opium poppy. As reported by
turmeric (Curcuma longa L.) was also successfully
Henry and Primo (1979), they observed a greater
enhanced by foliar application of TRIA (Singh 2008;
polyphenol oxidase activity in the leaf tissue of lettuce
Nasir 2009; Singh et al. 2011).
treated with foliar spray of TRIA in comparison to
the control. In an in vitro cell free system, the
activities of starch phosphorylase and phosphoenol
Role of TRIA in in vitro studies
pyruvate carboxylase (PEPCase) were increased up to
40% in the supernatant obtained from maize leaves, Generally, TRIA enhances the elongation of multiple
20 min after leaf treatment with TRIA (Houtz and shoots and micropropagated plantlets, although such
Ries 1983). an effect of TRIA has not been observed on the
Both in vitro and in vivo studies, conducted on rice woody plants (Tantos et al. 2001). The TRIA, applied
and maize, revealed a promotive effect of TRIA on at 10 mg L
1, increased the fresh and dry weight of
the content of reducing sugars, amino acids, and shoots by 20
25% in Salvia officinalis. The promotive
soluble protein in (Knowles and Ries 1981; Ries and effect of TRIA was also reported in micropropaga-
Wert 1982). Misra and Srivastava (1991) reported a tion of ornamentals and other plants (Kissimon et al.
significant effect of a TRIA formulation, Miraculan, 1999; Reddy et al. 2002). In an in vitro study,
on essential oil yield of lemongrass. They stated that Hangarter et al. (1978), demonstrated the favorable
Miraculan-spray at concentration of 0.4 mg L
1 effect of TRIA on Nicotiana tabacum, L. esculentum,

1.6 1200
(a) (b)
1.4 a a
a a
Artemisinin content (µg/g dw)

b a 1000
Essential oil content (%)

1.2 b
b
c c 800
1.0

0.8 600

0.6
400
0.4
200
0.2

0.0 0
Control T1 T2 T3 T4 Control T1 T2 T3 T4

Figure 3. Effect of foliar sprays of different concentrations of triacontanol (TRIA) and GA3 viz. Control, 1.5 mg TRIA (T1),
1.5 mg L
1 TRIA  50 mg L
1 GA3 (T2), 1.5 mg L
1 TRIA  75 mg L
1 GA3 (T3), and 1.5 mg L
1 TRIA  100 mg L
1 GA3
(T4) on essential oil content (a), artemisinin content (b) of Artemisia annua L. Bars showing the same letter(s) are not
significantly different at p  0.05 as determined by Duncan’s multiple range test. Error bars (-) show SE (Aftab et al. 2010).
 Journal of Plant Interactions 137

a
or 20 mg L
1) did not induce further increase in the
0.20
b diterpenoid yield of sage (Salvia officinalis) shoots.
bc bc
c Grzegorczyk et al. (2006) observed a positive effect of
TRIA on shoot multiplication, production of bio-
Essential oil content (%)

0.15
chemical compounds, and antioxidant capacity of S.
officinalis. Moreover, TRIA also increased carnosol
0.10 content in sage shoots, with a little influences on
carnosic acid. The ability of TRIA to increase
diterpenoid production may be ascribed to the
0.05
biosynthesis of the compounds that is probably
localized in the chloroplasts (Munne-Bosch and
0.00 Alegre 2001), the TRIA-mediated increase in chlor-
Control
–6 –6 –8 –6 –6
10 TRIA (T) 10 T+10 G 10 T+10 G 10 T+10 G
–6 –4
ophyll content in many plants being one of the strong
Moler concentrations evidences in this regard (Ries 1985).
Figure 4. Effect of spray of triacontanol (TRIA) alone or
in combination of GA3 on essential oil content of coriander.
Role of TRIA under abiotic stresses
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Means within a column followed by the same letter(s) are

not significantly different (p  0.05). The data shown are Muthuchelian et al. (2003) reported a TRIA-
means of four replicates 9 SE (Idrees et al. 2010). mediated increase in root and shoot, leaf density
and area, and fresh and dry biomass accumulation of
Solanum tuberosum, Phaseolus vulgaris, and a barley acidic-mist-treated Erythrina variegata plants. Their
hybrid (Hordeum vulgare H. jubatum). In addition, studies suggested that lipophilic TRIA might act on
encouraging results have been obtained with regard cell membranes to produce 9-b-L() adenosine. In
to TRIA-mediated growth improvement in O. sativa fact, this substance is rapidly translocated throughout
(Yun and Kim 1986), Malus domestica (Ma et al. the plant causing a cascade of metabolic events and,
1990), Melissa officinalis (Tantos et al. 1999), woody thus, resulting in significant increases in growth and
plants (Tantos et al. 2001), Bupleurum fruticosum dry matter of plant (Ries and Wert 1992).
(Fraternale et al. 2002), Capsicum frutescens and Application of TRIA increased 14CO2 fixation,
Decalepis hamiltonii (Reddy et al. 2002), and enzyme activities, synthesis of chl a, chl b, carote-
Thymus mastichina (Fraternale et al. 2003). Frater- noids, starch, and sugars in E. variegata seedlings
nale et al. (2003) recorded a TRIA-mediated enhance- under flooded and acid mist conditions as well as
ment in shoot growth of T. mastichina at 10 and 20 under salt and cadmium stressed conditions (Muthu-
mg L
1 of TRIA. Application of TRIA at a similar chelian et al. 1992, 1994, 1995, 1996, 1997, 2001,
range of concentrations produced improved number 2003). Muthuchelian et al. (2003) suggested that the
of nodes and shoot length as reported by others on TRIA-enhanced growth of plants might result from
mciropropagation of herbaceous and woody plants an increase in effective leaf area, stimulation of
(Tantos et al. 1999, 2001; Fraternale et al. 2002; photosynthesis and increase in the activities of
Reddy et al. 2002). Fraternale et al. (2003) reported RuBisCO and NR. Reductions in shoot growth
that TRIA applied at 10 and 20 mg L
1, enhanced the rate, leaf area index (LAI), and relative growth rate
essential oil yield of Thymus mastichina L. (about
3.5
33%) as compared to control. However, such an Total anthraquinone gycosides content
effect was not detectable when TRIA was applied 3.0 Total sennoside content

with auxin and/or cytokinin. Furthermore, they

2.5
observed that TRIA applied at 10 and 20 mg L
1 to
the control plants induced a decrease in the number 2.0
(%)

of glands at post-secreted stage (corresponding to an

1.5
increase of glands in secretory stage), and an increase
in the essential oil yield (about 67% increase in glands 1.0
in secretory-stage corresponding to about 47% in-
crease in the oil yield). 0.5

The biosynthesis of secondary metabolites in in 0.0

vitro culture studies is also affected by TRIA, like by 10–0 10–8 10–7 10–6 10–5
other plant growth regulators. TRIA has been TRIA concentrations (M)
reported to enhance the production of secondary Figure 5. Effect of five concentrations of triacontanol
metabolites in Artemisia annua (Yaseen and Tajuddin (TRIA) (10
0, 10
8, 10
7, 10
6, and 10
5 M) on total
1998). TRIA, applied at 5 mg L
1 enhanced the anthraquinone glycosides content and sennoside content
content of diterpenoids in the shoots of Salvia of coffee senna (Senna occidentalis L.) studied at 300 DAS
officianlis to the highest extent (Grzegorczyk et al. (Means of three replicates). Error bars (-) show SE (Naeem
2005). However, the use of TRIA concentrations (10 et al. 2010).
 138 M. Naeem et al.
6
100 100 DAP (a) 100 DAP (b)
120 DAP 120 DAP a
c c a a b b 5
c c

L-Menthone content (%)

b b
80 a

Menthol content (%)

4
60 c c
d c
3
40
2

20 1

0 0
10–0 10–7 10–6 10–5 10–0 10–7 10–6 10–5
TRIA concentrations (M) TRIA concentrations (M)

2.5 3.5
100 DAP (c) 100 DAP a a (d)
120 DAP 120 DAP b b

Menthyl acetate content (%)

3.0
a a
Isomenthone content (%)

2.0
d c c b b c c
d 2.5
d d
1.5 2.0
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1.5
1.0
1.0
0.5
0.5

0.0 0.0
10–0 10–7 10–6 10–5 10–0 10–7 10–6 10–5
TRIA concentrations (M) TRIA concentrations (M)

Figure 6. Effect of four concentrations of foliar sprays of triacontanol (TRIA) (10
0, 10
7, 10
6, and 10
5 M) on menthol
content (a), L-menthone content (b), isomenthone content (c), and menthyl acetate content (d) of mint (Mentha arvensis L.)
studied at 100 and 120 DAP. Means within a column followed by the same letter(s) are not significantly different (p  0.05).
Error bars (-) show SE (Naeem et al. 2011).

(RGR) were improved by TRIA application in water growth, yield, and physiological parameters in the
stressed seedlings of E. varietaga sprayed. Similar plants to a significant degree, with the exception of
results have been reported by Muthuchelian et al. electrolyte leakage, water saturation deficit and pro-
(1995) in E. variegata and by Srivastava and Sharma line content, whose values under these conditions was
(1991) in Papaver somniferum L. The reduction in the increased using TRIA. TRIA favorably influenced
contents of chlorophyll and carotenoids as well as both treated and untreated plants with periodic
that in chlorophyll fluorescence was also improved by chilling. Besides, a negative influence of chilling on
TRIA in salt-stressed E. variegata seedlings (Muthu- the growth and yield of Ocimum basilicum L. was
chelian et al. 1996) and Triticum aestivum plants. decreased by TRIA applied at 0.10 mg dm
3.
(Perveen et al. 2010). Thakur and Thakur (1993)
reported that application of TRIA and mixtalol
overcame the adverse effects of moisture stress in Further studies and future prospects
Lycopersicon esculentum cultivars. According to TRIA being a plant growth promoter plays pivotal
Muthuchian et al. (1994, 1997), TRIA application role in the up-regulation of many biochemical and
reduced the inhibition of PS-II activity to about 22% physiological processes in plants, including photo-
in water stressed plants of E. variegata; the reduction synthesis. This leads to maximize the growth, yield
in RuBisCO activity underwater stress was also and quality of several crops and helps improve the
improved by TRIA spray. They also reported a active constituents of medicinal and aromatic plants
TRIA-mediated maintenance in the photosynthetic under normal and abiotic stress conditions. The
machinery and a significant delay in the leaf-senes- application of TRIA may be successfully employed
cence in water stressed plants. By means of TRIA in future to enhance the productivity, quality, and
application, Krishnan and Kumari (2008) reported a production of essential oil and other active constitu-
successful amelioration of salt stress in soybean ents of medicinal and aromatic plants. However,
plants in terms of leaf weight ratio, relative water further investigations are required to elucidate the
content, chlorophyll pigments, nucleic acids, soluble possible regulatory role of TRIA on plant growth,
sugars, and soluble proteins. physiological activities, and biosynthesis of secondary
By TRIA application, Rajasekaran and Blake metabolite in medicinal and aromatic plants subjected
(1999) reversed the damaging effect of drought-stress to abiotic stresses. Intense efforts are still required in
in terms of membrane leakage on Jack pine seedlings. order to design the protocols to ensure the consistent
The study of Borowski and Blamowski (2009) showed results regarding response of plants to application of
that periodic chilling decreased the value of all the TRIA and l()-adenosine in the growth chambers,
 Journal of Plant Interactions 139

greenhouses, hydroponics systems, and field studies. Asane GB, Lawande KE, Nirmal SV, Shinde KG, Desale
It is also a significant challenge to discover as to how SB. 1998. Effect of cytozyme crop, triacontanol and
both TRIA and l()-adenosine rapidly increase the cycocel on growth, yield and quality of pea (Pisum
plant metabolism and how TRIA elicits the synthesis sativum L.). Advan Plant Sci 11:31
34.
Barus SC. 1998. Productivity of Santa Rosa plum trees in
of l()-adenosine. Identification of OCTAM and
response to triacontanol. J Inter Academician 2:
determining as to how it inhibits the activity of TRIA
124
129.
may lead to the discovery of the site of TRIA and/or Bhalla RP. 1981. Triacontanol as plant biostimulant.
l()-adenosine action (Ries et al. 1993). Proceedings 8th annual meeting of the plant growth
Since the PGRs that boost up crop productivity regulator Society of America. August 3
6; St. Peters-
are normally expensive, it is always desirable to find burg Beach, Florida, p. 184
195.
out the low-cost PGRs. The TRIA fulfills this Blamowski ZK, Borowski E, Blamowska M. 1998. Wplyw
requirement as it is extraordinarily cheap to afford. dtugotaficuchowych alkoholi alifatycznych na wzrost,
In future studies, it could also be explored if TRIA, in wymian  gazow  i rozdzial asymilat6w w roglinach
combination with other inexpensive PGRs, could rzodkiewki. Acta Agrobot 51:5
10.
further enhance the performance of crop plants at a Borowski E, Blamowski ZK. 2009. The effects of triacon-
desirable extent. The secondary messengers, which tanol ‘TRIA’ and Asahi SL on the development and
metabolic activity of sweet basil (Ocimum basilicum L.)
are produced by TRIA and other growth promoting
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plants treated with chilling. Acta Hort 21/1:39
48.

substances, and the mechanism of TRIA-mediated
Borowski E, Blamowski ZK, Michalek W. 2000. Effects of
elicitation of plants metabolism could also be worked tomatex/triacontanol on chlorophyll fluorescence and
out in future studies. In addition, the research work tomato (Lycopersicon esculentum Mill.) yields. Acta
regarding combined application of TRIA and other Physiol Plant 22:271
274.
PGRs would enable one to explore the economically Bouwkamp JC, McArdle RN. 1980. Effects of triacontanol
and ergonomically feasible agronomic solutions to on sweet potatoes. Hort Sci 15:69.
increase the essential oil and the medicinally impor- Chaudhary BR, Sharma MD, Shakya SM, Gautam DM.
tant active constituents in medicinal and aromatic 2006. Effect of plant growth regulators on growth,
plants, the production of which is far short than it is yield and quality of chilli (Capsicum annum L.) at
unanticipated high demand in the global market. Rampur, Chitwan. J Inst Agric Animal Sci 27:65
68.
Chen MZ, Kuo CH, Dau S, Sau TL. 1980. Physiological
effects of 1-triacontanol on winter wheat. Rpt Sci
Conclusions Technol J 12:1
3.
Chen X, Yuan H, Chen R, Zhu L, Du B, Weng Q, He G.
The evidence is overwhelming that TRIA applied at 2002. Isolation and characterization of triacontanol-
nanomolar concentrations improves the plant growth regulated genes in rice (Oryza sativa L.): Possible role
and physiological activities in diverse groups of of triacontanol as a plant growth stimulator. Plant Cell
plants. Foliar application of TRIA has been proved Physiol. 43:869
876.
to be a successful technique to improve the growth, Chen X, Yuan H, Chen R, Zhu L, He G 2003. Biochemical
yield, and quality of various crops, including vege- and photochemical changes in response to triacontanol
tables, horticultural crops, and medicinal and in rice (Oryza sativa L.). Plant Growth Regul 40:
aromatic plants. TRIA-mediated increase in dry 249
256.
matter leads to increased biosynthesis of secondary Chowdhury SR, Anand PSB, Ashwani K. 2009. Triaconta-
plant products as well, including essential oil and nol induced changes in kernel dry matter, carbohy-
drate content and yield of water chestnut (Trapa
active constituents of medicinal and aromatic plants.
bispinosa L.) fruit. Indian J Plant Physiol 14:88
92.
However, further studies are required to reveal the
Diehi H. 1974. Quantitative analysis. 2nd ed. Ames (IA):
possible role of TRIA on the regulation of plant Oakland Street Press. p. 173
177.
growth and metabolism in terms of regulation of gene Eriksen AB, Haugstad MK, Nilsen S. 1982. Yield of
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applications of triacontanol. Plant Growth Regul
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14.
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Financial support by the Science and Engineering Research Comparative analysis of the effect of triacontanol on
Council, Department of Science & Technology, Govern- photosynthesis, photorespiration and growth of toma-
ment of India, New Delhi, for the award of Young Scientist to (C3-plant) and maize (C4-plant). Planta 152:44
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to Dr. M. Naeem (Project No. SR/FT/LS-003/2008) is Farooqi AHA, Sharma S 1988. Effect of growth retardants
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