Puncturevine (Tribulus terrestris L.): Noxious weed or powerful medical herb

Abstract

Tribulus terrestris L., an annual dicot species of the family Zygophyllaceae, is a common herb that is often found in disturbed habitats and agricultural areas in many parts of the temperate, tropical and desert regions of the world. T. terrestris is an aggressive species that has the potential to injure livestock, reduce hay and wool values, detour recreationists and reduces plant biodivesity. The species may become troublesome because of its weedy potential. It has been declared a weed in at least 37 countries and in at least 21 crops (cotton, maize, vineyards, orchards, etc.). It is adapted to a wide range of climatic conditions and grows on a wide variety of soil types. The management of T. terrestris can be achieved by herbicide application, mechanical (hand pulling, hoeing, mulching) and biological control methods. Beside its invasive potential as a noxious and troublesome weed, T. terrestris is considered highly useful herb which is used for various purposes in folk and modern medicine and sport, as well.

Full text

Puncturevine (Tribulus terrestris L.): noxious weed
or powerful medical herb
Kotvičník zemný (Tribulus terrestris L.): nebezpečná
burina alebo silná liečivá rastlina
Zvonko PACANOSKI1, Štefan TÝR2* and Tomáš VEREŠ2
1 Ss. Cyril and Methodius University in Skopje, Faculty for Agricultural Sciences and Food, blvd.
Aleksandar Makedonski bb 1000, Skopje, Republic of Macedonia, zvonko_lav@yahoo.com
2 Slovak University of Agriculture in Nitra, Faculty of Agrobiology and Food Resources, Nitra, Tr. A.
Hlinku 2, 949 76 Nitra, Slovakia, Stefan.Tyr@uniag.sk*correspondence
Abstract
Tribulus terrestris L., an annual dicot species of the family Zygophyllaceae, is
a common herb that is often found in disturbed habitats and agricultural areas in
many parts of the temperate, tropical and desert regions of the world. T. terrestris is
an aggressive species that has the potential to injure livestock, reduce hay and wool
values, detour recreationists and reduces plant biodivesity. The species may become
troublesome because of its weedy potential. It has been declared a weed in at least
37 countries and in at least 21 crops (cotton, maize, vineyards, orchards, etc.). It is
adapted to a wide range of climatic conditions and grows on a wide variety of soil
types. The management of T. terrestris can be achieved by herbicide application,
mechanical (hand pulling, hoeing, mulching) and biological control methods. Beside
its invasive potential as a noxious and troublesome weed, T. terrestris is considered
highly useful herb which is used for various purposes in folk and modern medicine
and sport, as well.
Keywords: agriculture and medical importance, biology, control, ecology, Tribulus
terrestris
Abstrakt
Kotvičník zemný (Tribulus terrestris L.), jednoročný dvojklíčnolistový druh z čeľade
jarmovcovité (Zygophyllaceae), je bežnou liečivou bylinou, ktorá sa často vyskytuje
na narušených stanovištiach a na poľnohospodárskej pôde v mnohých častiach
teplých, tropických a púštnych oblastí sveta. T. terrestris je agresívny druh, ktorý
potenciálne môže poraniť hospodárske zvieratá, redukovať hodnotu sena či vlny
a redukuje diverzitu rastlinného krytu. Tento rastlinný druh môže spôsobovať problem
pre jeho potenciál stať sa burinným druhom. Bol deklarovaný ako burina v najmenej
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Journal of Central European Agriculture, 2014, 15(1), p.11-23
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Journal of Central European Agriculture, 2014, 15(1), p.11-23 DOI: 10.5513/JCEA01/15.1.1404

37 krajinách a v najmenej 21 plodinách (bavlna, kukurica, vinohrady, sady, ai.). Je
adaptibilný na široký rozsah klimatických podmienok a rastie na rozmanitých
pôdnych typoch. Manažment regulácie T. terrestris zahŕňa aplikáciu herbicídov,
mechanickú (ručné vytrhávanie, bránenie, nastielanie) a biologickú reguláciu.
Napriek jeho inváznemu potenciálu ako nebezpečná a problémová burina,
T. terrestris je považovaný za vysoko užitočnú liečivú rastlinu, ktorá je používaná na
rôzne účely v ľudovej aj modernej medicíne a v športe.
Kľúčové slová: poľnohospodársky a medicínsky význam, biológia, regulácia,
ekológia, kotvičník zemný, Tribulus terrestris
Detailný abstrakt
Kotvičník zemný (Tribulus terrestris L.) je jednoročná dvojklíčnolistová bylina z
čeľade jarmovcovité (Zygophyllaceae) pochádzajúca zo Saharskej oblasti. Je bežnou
liečivou bylinou, ktorá sa často vyskytuje na narušených stanovištiach a na
poľnohospodárskej pôde v mnohých častiach teplých, tropických a púštnych oblastí
sveta. Na druhej strane T. terrestris je agresívny druh, ktorý potenciálne môže
poraniť hospodárske zvieratá, redukovať hodnotu sena či vlny a taktiež redukuje
diverzitu rastlinného krytu. Tento rastlinný druh môže spôsobovať problem pre jeho
potenciál stať sa burinným druhom. Bol deklarovaný ako burina v najmenej 37
krajinách a v najmenej 21 plodinách (bavlna, kukurica, vinohrady, sady). Tribulus
terrestris sa stáva burinným druhom aj kvôli svojmu hlboko rozkonárenému
drevnatému koreňu. Rastlina vytvára veľké množstvo bočných vetiev dlhých až 3 m a
tým pokrýva pôdu a vytvára hustý porast. Je adaptibilný na široký rozsah
klimatických podmienok a rastie na rozmanitých pôdnych typoch. Manažment
regulácie T. terrestris zahŕňa aplikáciu herbicídov, mechanickú (ručné vytrhávanie,
bránenie, nastielanie) a biologickú reguláciu. Najdôležitejším opatrením pri regulácií
kotvičníku zemného je regulácia jeho generatívnych orgánov rozmnožovania
v pôdnej zásobe a zabránenie v tvorbe semien a plodov všetkými dostupnými
prostriedkami. Napriek jeho inváznemu potenciálu ako nebezpečná a problémová
burina, T. terrestris je považovaný za vysoko užitočnú liečivú rastlinu, ktorá je
používaná na rôzne účely v ľudovej aj modernej medicíne a v športe.
Introduction
Tribulus terrestris L. (puncturevine, punctureweed, goatshead), is an herbaceous,
monocarpic, a C4 summer annual, broadleaf weed a member of the Zygophyllaceae
or caltrop family (Donaldson and Rafferty, 2003; Šalamoun et al., 2006). It is occurs
widely throughout the world from latitudes 35°S to 47°N (Holm et al., 1991). T.
terrestris is native to southern Europe (Grin, 2000, Parker 1972), Africa, temperate
and tropical Asia, and north Australia (Grin, 2000). According Squires, (1979) T.
terrestris probably originated in the Saharan region, and spread into the
Mediterranean region. It is one of the most widely distributed species and well
adapted to temperate, mediterranean, sub tropical, tropical and warm temperate
climate (Lamp and Collet, 1990; Scott and Morrison, 1996). T. terrestris requires
relatively high temperatures for growth and is prevalent in areas having hot summers
(WSNWCB, 2001; CDFA, 2002), but it is intolerant of freezing temperatures (Squires,
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1979). Boydston (1990) notes that T. terrestris can be found in a wide range of
conditions: it thrives on dry, loose, sandy soils and prospers near sand dunes and in
wind-blown loose soil by field margins; however, it also grows in heavier soils,
especially if they are fertile and moist, and on compacted soils such as those found
along the sides of unsurfaced roads or in playgrounds (El-Ghareeb, 1991). T.
terrestris is an aggressive species that has the potential to injure livestock
(Glasonbury et al., 1984; Kellerman et al., 1994; McDonough, 1994), reduce hay and
wool values (Gould & Deloach, 2002; Knight and Walter, 2003; SIR and EPD, 2004)
and detour recreationists (Donaldson and Rafferty, 2003). T. terrestris reduces plant
biodivesity by quickly invading and crowding out desirable species (Van Vleet, 2005).
The species may become troublesome because of its weedy potential (Boydston,
1990; Scott and Morrison, 1996; Geier and Stahlman, 1999). Due to its ability to
extract soil moisture from great depth in the soil, T. terrestris competes well in many
crops (Holm et al., 1991). It has been declared a weed in at least 37 countries and in
at least 21 crops (cotton, maize, vineyards, orchards, etc.) (Asher et al., 2002; Kostov
& Pacanoski, 2007; Verd„u and Mas, 2007; Cheema et al., 2008; Geier et al., 2006;
Tahir et al., 2009; Kir & Dogan, 2009). Out of cultivated fields, T. terrestris exist on
disturbed places, along streets, roadsides, railways, waste places, walk ways,
pastures, lawns and yards, etc. (Hickman, 1993; Guertin and Halvorson, 2003;
CDFA, 2002). Beside its invasive potential as a noxious and troublesome weed, T.
terrestris is considered highly useful herb. It is an herbal remedy which is used for
various purposes in folk medicine. Ancient Greeks used T. terrestris as a diuretic and
a mood-enhancer. In ancient Chinese medicine, it was used for a variety of liver,
kidney, and cardiovascular diseases (Sahelian, 2003). Traditional herbs have
emerged in the past few years as an „instant‟ treatment for sexual and erectile
dysfunctions (Adimoelja, 2000).
Taking into consideration previous mentioned facts, Tribulus terrestris is common in
many parts of the world and it has remarkable features as a cosmopolitan weed and
highly invasive and aggressive species. Thus, the aim of this report was to
summarize the available information and bring together new information and recent
trends particularly from an agronomic point of view i.e. agricultural importance and
management of puncturevine Tribulus terrestris.
Biology and ecology of Tribulus terrestris
Tribulus terrestris has a deep woody taproot. The plant produces numerous prostrate
stems, up to 3 m long that are much branched and arise from the crown to produce a
dense mat. The leaves are cotyledons oblong, opposite, short-petioled, 2-5 cm long,
pubescent, and divided into pinnate elliptic or oblong leaflets (3-7 leaflet pairs per
leaf); each leaflet 3-15 mm long. The small, yellow, 5-petaled perfect flowers are
borne on short stalks at leaf nodes. The fruit is a schizocarp; woody burr with sharp,
rigid spines to approximately 1-1.8 cm in diameter. Seeds are usually 2-5 per burr,
and remain enclosed within the burrs (Yingxin, 1998; Donaldson and Rafferty, 2003;
Kostov, 2006; WSNWCB, 2008). A plant may produce 200 to 5,600 seeds during one
growing season (Boydston, 1990), and a large plant may produce up to 10,000 seeds
(CDFA, 2002). Various studies showed seeds can remain viable for several years
(CDFA, 2002), staying dormant in the soil for 4-5 years (Whitson, 1992). Humans
and their activities as well as animals are the most important means of seed
dispersal (Ernst and Tolsma, 1988; Squires, 1979; Whitson 1992).
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Seeds germinate from spring to autumn under suitable moist and warm conditions
(from 24 °C to 27 °C) and it grows rapidly producing deep root system in a few
weeks. (Scott and Morrison, 1996; CDFA, 2002). Ernast and Tolsma (1988)
observed that germination of T. terrestris in the field can start after a rain shower of
more than 10 mm. T. terrestris can flower within 3–4 weeks after emergence when
temperatures are above 20 °C (Boydston, 1990) primarily from July to August
(Parker, 1972). Once the plant begins to flower, it is continuous throughout the plant's
life (Reddi et al., 1981). Fruits mature in approximately 2 weeks, and subsequently
split apart into segments soon afterward (Holm et al., 1991). Plants continue to
reproduce and produce fruit until the cool season begins. Boydston (1990) reports
during trials in Washington, fruit/burr production stopped in October when average
temperatures were under 20°C. Seeds can be produced as soon as 5-6 weeks after
germination (Scott and Morrison, 1996; CDFA, 2002; WSNWCB, 2001). Because of
its large seed production and the long-term viability of seeds, this species can
increase in numbers rapidly under suitable conditions (Boydston, 1990). The plants
usually die in autumn or sinter after the first frosts (Squires, 1979). Generally, T.
terrestris has a considerable seed dormancy lasting over fall and winter months
(WSNWCB, 2001) with some seeds staying dormant for longer periods of time, but in
tropical areas it was observed that seeds may still germinate in the fall (Pathak,
1970). In these areas under suitable conditions T. terrestris develops woody roots
and becomes perennial (Holm et al., 1991; CDFA, 2002).
Tribulus terrestris is adapted to a wide range of climatic conditions. It is prevalent in
areas having hot summers (Boydston, 1990) in warm, temperate and desert regions
(WSNWCB, 2001). T. terrestris requires relatively high temperatures for growth, and
is intolerant of freezing temperatures (CDFA, 2002; Squires, 1979). It occurs in areas
with a mean annual minimum precipitation of 280 mm and a mean annual maximum
precipitation of 380 mm (Rice, 2002). Seedling establishment was observed to be
poor on sites that were shaded (Pathak, 1970). T. terrestris grows on a wide variety
of soil types, but it is found most commonly on dry, loose, sandy soils and prospers
near sand dunes and in wind-blown loose soil by field margins (WSNWCB, 2001;
CDFA, 2002); however, it also grows in heavier soils, especially if they are fertile and
moist (Holm et al., 1991), and on compacted soils such as those found along the
sides of unsurfaced roads or in playgrounds (El-Ghareeb, 1991).
Agricultural importance of Tribulus terrestris
Tribulus terrestris is considered to be an aggressive, highly invasive species,
problematic and “cosmopolitan” weed with a worldwide distribution, that is mainly
attributed to human activity (Van Vleet, 2005). T. terrestris requires disturbance to
establish and is most often associated with an anthropogenic disturbance. Because
the ecological amplitude of T. terrestris is so broad, it can invade most ecological
types in Arizona when they are anthropogenically disturbed to a significant degree
(Guertin, 2001). Puncturevine is classified as a class B designate noxious weed in
Washington and among invasive plant species with the greatest and most immediate
threats to the biological resources (Evans et al., 2003).
In cropping systems, Tribulus terrestris decreases crop yield through competition for
sunlight, soil water, and nutrients. In Pakistan a list of important weed species of
cotton crop includes T. terrestris (Hakoomat et al., 2005) with frequency of 53.75 %
(Memon et al., 2007). T. terrestris with Cyperus rotundus are the most dominant
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weeds in maize fields in Pakistan (Tahir et al., 2009). According to Marwat, (1984)
and Ahmad et al. (2000) T. terrestris is one of the most serious weeds that cause
damage to the maize and soybean crop in this country. A phytosociological study of
weeds carried out at Ajmer in the agriculture semi-arid zone of India, revealed the
dominant presence of Cynodon-Tribulus-Tephrosia community of weeds in many
crops during the summer season, growing on coarse and sandy soils (Sharma,
1981). In accordance with these results, T. terrestris is recorded as a problematic
weed in soybean (Singh and Jolly, 2004), sugarcane (Singh and Kaur, 2003),
chickpea (Aujla and Cheema, 1983), onion (Randhawa and Bhalla, 1976) and young
peach orchards (Chatha and Chanana, 2007). T. terrestris (37.5 %) was detected as
a major weed species together with Amaranthus spp. (41.5 %), Sorghum halepense
(29%), Echinochloa crus-galli (25 %) and Convolvulus arvense (24%) in dry bean
(Ahmadi et al., 2007), and, also in maize fields in Iran (Mahmoodi and Rahimi, 2009).
Results of Kir and Dogan, (2009) and Uremis et al. (2004) showed that T. terrestris is
a weed with high frequencies in maize fields and one of the most common weeds in
strawberry growing areas (Boz, 2003) in Turkey. T. terrestris is one of the most
common weeds of cotton (Chenault et al., 1986; Asher et al., 2002) and green bean
(Black et al., 2003) in Texas, as well, but weed with less density in sorghum (Wiese
et al., 1964; Hennigh et al., 2010) and maize in Kansas (Geier and Stahlman, 1999;
Geier et al., 2006). In Republic of Macedonia, T. terrestris is important weed with
negative economical impact on melon crops (Kostov, 2006; Kostov and Pacanoski,
2007). Also, T. terrestris was recorded as a troublesome weed in the peanuts in
Cyprus (Vouzounis, 2006), tomato in Southern Italy (Tei et al., 2003), capsicums and
chillies in Australia (Frost and Hingston, 2006) potato in Sudan (Mohamed and Nour,
1986), safflower in Colorado (Anderson, 1985) and mandarin orchards in Spain
(Verd´u and Mas, 2007). Johnson and Talbert (1993) reported that T. terrestris is a
problematic weed in peas used by commercial vegetable processors, because its
burrs and seeds can be difficult to remove from peas and lima bean seed (Parker
and Boydston, 2007). T. terrestris is a nuisance weed in alfalfa beacause
contaminated hay can contain high levels of nitrates and burs can injure mouths of
livestock, lowering the value and quality of the hay (Boydston, 2010). The stiff, sharp
spines are a nuisance in many settings and grazing of the foliage can poison
livestock (Squires, 1979; Kostov, 2006). Sheep eating T. terrestris develop
photosensitivity secondarily to biliary obstruction that is result of steroidal saponins in
the plant (Glasonbury et al., 1984; Knight and Walter, 2003). Severe effects include
blindness, necrosis of skin, loss of lips and ears, and death in young animals.
Integrated Management of Tribulus terrestris
Taking into consideration fact that Tribulus teresterris is aggressive, highly invasive
and toxic species, problematic and “cosmopolitan” weed, control methods should be
combined into an integrated management system for the best long-term control of
this weed. Management techniques selected are dependent upon a specific site and
will be determined by land use objectives, extent of T. terrestris infestations and
effectiveness and limitations of available control measures (Schultz, 2005). Long-
term control of T. terrestris can be achieved by reducing the amount of seeds in the
soil. This is best accomplished by removing plants before they produce seeds (i.e.
before or at flowering) and continuing to do so over several years (Donaldson and
Rafferty, 2003).
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A number of herbicides are effective in various crops and situations against T.
terrestris, but its control is difficult because seeds can germinate throughout summer
and then rapidly flower and produce viable seed (Affeldt and Campbell, 2007) which
may enable T. terrestris to persist in spite of weed control programs (Boydston,
1990). Guethle et al., (1990) found imazethapyr at 0.07 kg a.i.*ha-1 applied PPI
controlled 92 % of T. terrestris and reduced its seed 94%Imazethapyr applied to
southern peas at 0.07 kg a.i.*ha-1 PPI, PRE, and 3 and 6 d after T. terrestris
emergence controlled T. terrestris at least 95% at the 2-week rating. At the 4-week
rating, imazethapyr at 0.07 kg a.i.*ha-1 applied PRE provided this level of control.
Imazaquin at 0.07 and 0.14 kg a.i.*ha-1 controlled T. terrestris greater than 93% when
applied up to 12 d after emergence (Johnson and Talbert, 1993). Successful control
of T. terrestris and other weeds in peanuts was achieved through the pre-and post-
emergence application of imazethapyr at 0.1 to 0.12 kg a.i.*ha-1 (Vouzounis, 2006).
The results of Tahir et al., (2009) showed that pendimethalin applied at 1,050 g
a.i.*ha-1 and pendimethalin + prometryn applied at 1,400 g a.i.*ha-1 significantly
controlled (78.25%, and 69.57%, respectively) T. terrestris in maize over the weedy
check. Based on the ED90 values D. stramonium, T. terrestris, E. crus-galli, S.
halepense, A. retroflexus, A. blitoides, and S. nigrum were highly sensitive to
foramsulfuron and were controlled with less than 50% of the recommended herbicide
rate in maize (Kir and Dogan, 2009). Control of T. terrestris in corn with EXP 31130A
alone or in tank mixtures with acetochlor, atrazine and metolachlor was 75 % or
greater (Geier and Stahlman, 1999). In investigation of Geir et al., (2006) T. terrestris
control in corn exceeded 94 % with KIH-485 and S-metolachlor applied at different
rates, but mixtures of atrazine with KIH-485 or S-metolachlor generally provided the
most effective control of T. terrestris and other broadleaf weeds studied. Oxadiargyl
at 400 g a.i.*ha-1 was generally more effective than clomazone at 480 g a.i.*ha-1 and
pendimethalin at 660 g a.i.*ha-1 for controlling T. terrestris in capsicum and chillies
(Frost and Hingston, 2006). Pendimethalin (1.48 kg a.i.*ha-1) and oxidiazon (0.45 kg
a.i.*ha-1) showed excellent performance in controlling of T. terrestris and other weeds
in autumn sown soybean (Ahmad et al., 2000). In alfalfa, preemergence applied
flumioxazin and norflurazon control early season T. terrestris germination. Imazamox,
2,4-DB, and bromoxynil applied postemergence control T. terrestris seedlings less
than 4 cm tall (Boydston, 2010). Glyphosate plus metsulfuron and glyphosate plus
2,4-D ester gave an average of 90 and 88% control, respectively of the T. terrestris
and other summer-growing weeds on fallows in southern Australia (Leys et al.,
1990).
Tribulus teresterris can be managed using mechanical (hand pulling, hoeing, and
mulching) and biological control methods, as well. As with all annuals, mechanical
controls are partially effective in control of T. terrestris. On small infestations, hand-
pulling prior to flower and seed production is effective in controlling new infestations
(CNAP, 2000), but mowing is not effective because of the low growth habit of the
plant. Hoeing and shallow cultivation (about 2-3 cm deep) are, also effective at killing
existing plants, and should be initiated prior to flowering and seed production
(Fenner, 1985; Holm et al., 1991; Donaldson and Rafferty, 2003). If plants have
produced fruits before a cultivation effort is made, subsequent to the effort the plants
and fruits should be collected and burned (Muenscher, 1980; WSNWCB, 2001).
Several years‟ cultivation may be required to exhaust the seedbank in established
infestations (WSNWCB, 2003). Mulches can be used to control T. terrestris in
ornamental plantings, orchards, vineyards, vegetable crops, and gardens, if they
screen out all light. Results of Verd´u and Mas, (2007) indicated that black geotextile
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and almond husk, used as mulches, controlled the presence of 74 weeds species in
mandarin orchards, including T. terrestris, as well as or better than the applications of
glyphosate at least during the first year after their introduction. Grazing as a control
method is not allowed, because plant is toxic and can cause physical injury to
animals (Holm et al., 1991).
Biological control agents have been relatively successful for Tribulus terrestris
control. It is controlled by two weevils native to India, France, and Italy (WSNWCB,
2003). They are the stem weevil (Microlarinus lypriformis) and the seed weevil
(Microlarinus lareynii) (Hickman, 1993). The larvae attack the stems and seed and
have reportedly provided reasonably good results (WSNWCB, 2003). Both insects
provide good control of the plant, but it may take several years to deplete the seed
bank in the soil. Good biological control has been achieved in Hawaii (Julien and
Griffiths, 1998) where T. terrestris being completely eradicated within 4 years (Markin
et al., 1992). Wilson et al., (1997) and Gould and Deloach (2002) reported for partial
success in biological control of T. terrestris in some areas of Nevada, California,
Arizona, Texas and New Mexico. These insects have not overwintered in many
northern latitudes.
Medical uses and values of Tribulus terrestris
Tribulus terrestris is a strong herbal remedy which is used for various purposes in
folk and modern medicine and sport, as well. It has been used as tonic, aphrodisiac,
astringent, analgesic, stomachic, anti-hypertensive, antibacterial, antifungal and
urinary anti-septic (Kianbakht and Jahaniani, 2003; Al-Bayati and Al-Mola, 2008).
According to Arcasoy et al. (1998), T. terrestris has been commonly used as a
diuretic as well as treatment for hypertension, hypercholesterolemia and colic pains.
Wang et al. (1990) found that T. terrestris supplementation may reduce the remission
rate of angina pectoris and decrease myocardial ischemia without any unwanted
effects on hepatic or renal function. Dimitrov et al., (1987) found increased plasma
testosterone levels and reversed sexual impotence in rams following
supplementation with T. terrestris. It has a complex stimulating effect on germinative
and endocrine functions of the testes producing its precocious development (Bashir
et al., 2009). Arsyad (1996) showed that T. terrestris (protodioscin) treatment led to
an invariable increase in concentration of spermatozoa in humans to approximately
160%. The author attributed this to an increase in the LH (luteinizing hormone) level
which acted on Leydig cells and enhanced testosterone secretion, and stimulated
Sertoli and germinal cells. T. terrestris is considered an aphrodisiac, a putative
testosterone elevator. It increase sexual function in animal studies and also
reportedly improves libido in humans (Adaikan et al. 2000, Dimitrov et al. 1987). T.
terrestris has been used for centuries in Europe as treatment for impotence (Sharifi
et al. 2003). It enhances plasma testosterone levels and promotes skeletal muscle
hypertrophy. Supplement manufacturers claim that T. terrestris enhances
testosterone production via the stimulation of luteinizing hormone from the pituitary
glands; thus, gain in skeletal muscle mass may occur secondary to an augmentation
of plasma testosterone. Because of that, T. terrestris was and still is a source of the
success and top secret of many sport stars in the past and nowadays.
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