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CFE also increased the pro-collagen type-I N-terminal propeptide: cross-linked C-telopeptide of type-I collagen PINP : CTX-1 ratio over the OVX rats, and maintained it to the sham level.
CFE treatment decreased the OVX-induced increases in the carbonate-to-phosphate, and carbonate-to-amide-I ratios. CFE also prevented the OVX-mediated decrease in mineral crystallinity. Nanoindentation parameters, including modulus and hardness, were decreased by OVX but CFE maintained these to the sham levels.
Forskolin stimulated ALP, cAMP and cGMP in vitro and upregulated osteogenic genes in vivo. Conclusion: CFE, likely due to the presence of forskolin displayed a bone-conserving effect via osteogenic and anti-resorptive mechanisms resulting in the maintenance of bone mass, microarchitecture, material, and strength.
Coleus forskohlii CF , commonly known as Coleus in English, is a medicinal herb with rich ethnopharmacological applications. CF is also used in Ayurvedic medicine to treat a variety of ailments, including inflammatory diseases, hypertension, respiratory disorders, aging, and weight management 1 , 2.
CF is a rich source of secondary metabolites, including terpenoids, flavonoids, and alkaloids 3. The major bioactive compound of Coleus root is forskolin, a labdane diterpene which is of clinical interest because of its weight-loss property.
CF is the only species known to contain significant amount of forskolin 4. Forskolin acts by increasing the accumulation of cyclic adenosine monophosphate cAMP without hormonal stimulation of adenylate cyclase AC 2 , 5. cAMP binds to, and activates protein kinase A PKA , which then activates lipases by phosphorylating them, resulting in lipolysis.
Given this mechanism of action of forskolin, CF extract is thought to have an anti-obesity effect that has been demonstrated in a few preclinical studies 5 — 8.
Human studies, albeit scanty present inconsistent outcomes in reducing the weight of obese men and women 5 , 9 , 10 although it could attenuate weight gain Activation of AC by forskolin resulting in the rise in intracellular cAMP appears to underlie the osteogenic effect of the compound.
The osteogenic drugs teriparatide PTH and abaloparatide PTHrP act by type 1 PTH receptor to activate AC to increase intracellular cAMP in osteoblasts However, cAMP can either stimulate or inhibit osteogenic differentiation in human mesenchymal stem cells, depending on the duration of the rise in its intracellular levels Rises in the intracellular cAMP stimulate mesenchymal stem cells MSC proliferation and differentiation to osteoblasts, as well as osteoblastic differentiation from pre-osteoblasts db-cAMP, a synthetic cAMP analog, stimulated osteogenic differentiation in vitro and new bone formation in vivo Sustained stimulation of cAMP signaling, on the other hand, decreases osteoblast differentiation and mineralization 15 while inducing adipocyte differentiation We previously observed that the profile of cAMP activation kinetics of forskolin matches with PTH 15 , which led us to surmise that CF rich in forskolin may have an osteogenic property.
Here, we used a standardized preparation of CF root extract CFE, rich in forskolin and studied the osteogenic and anti-osteoporotic effects in rats.
Three models were used for these purposes, i femur osteotomy model for rapid assessment of bone regeneration for determining the osteogenic dose of CFE, ii growing rats for determining the modeling-directed bone formation in intact rat bones, and iii OVX rats a model for postmenopausal osteopenia to assess the effect of CFE on maintaining bone mass, microarchitecture, bone formation, bone turnover, bone strength, and bone quality.
Ex vivo cultures of bone marrow cells were used to evaluate the effect of CFE on osteoblast differentiation. Finally, we determined the amount of forskolin in CFE and assessed its in vivo osteogenic effect.
CFE used in this study was procured from Pharmanza Herbal Pvt. Ltd Anand, Gujrat, India. Forskolin was procured from Phytocompounds Bangalore, India.
Acetonitrile and methanol MS grade were procured from JT Baker and Rankem. Cell culture medium and all chemicals were procured from Sigma-Aldrich St. Louis, MO, USA. FBS, collagenase and diaspase were purchased from Invitrogen Carlsbad, CA, USA.
Gum acacia was purchased from Santa Cruz Biotechnology, Inc. Dallas, TX, USA. The solution of forskolin standard 1. The mixture was then centrifuged, filtered, and used for further analysis using HPLC according to our previously described method The analysis was performed on a Phenomenex ® Luna C18 column ×4.
With gradient elution of water and acetonitrile at a flow rate of 1. The column temperature was kept at 30°C, with an injection volume was Animal husbandry and all animal experimental procedures were prior approved by the Institutional Animal Ethics Committee Registration no.
The rats were acclimated for 8 days prior to surgery. The rats were maintained on standard rodent chow diet and purified water ad libitum during the experimental period.
Adult rats with a drill-hole 0. Furthermore, this model is useful for the determination of effective osteogenic dose of a drug. Twenty four female SD rats ± 20 g were used for femur osteotomy following a previously described protocol All the treatments were given daily for 12 days.
After sacrifice, bones were collected and processed for calcein labeling studies according to our previously published protocol. Sections were photographed using a confocal microscope Leica TCS SP-8, Wetzlar, Germany and analyzed using LAS-X software.
Modeling-directed bone formation is the dominant event in growing rats, which enables evaluation of the osteogenic response in intact skeleton 21 , Treatments were given for 1 month. For dynamic histomorphometry study to measure surface-referent bone formation parameters , each animal was given two s.
At the end of the experiment, bones were collected for μCT, bone strength, and histomorphometry analyses, for which the details are given below.
Femur length was measured with a Vernier caliper. To this aim, 24 SD rats ± 20 g, 3 months old underwent either bilateral OVX or a sham operation ovary intact 19 , All the therapies were given daily for 3 months.
before sacrifice at the interval of 10 days 19 , After the treatment, serum samples, and bones femur, tibia, and L5 vertebrae were dissected and stored at °C for further studies.
Throughout the experimental period, the body weights of each study group were measured once a week. Singapore 24 h before the end of the experiment Total body mass and lean mass were plotted, while fat mass was plotted by normalizing with the total body mass.
Bone samples were scanned using a SkyScan computed tomography μCT scanner SkyScan, Ltd. CTAn software was used manually to quantify various bone parameters as described previously Reconstructed μCT images underwent a blind evaluation by a third person to determine the extent of bone loss.
The degree of bone loss was assessed using reconstructed μCT images that had undergone a blind examination by a third person. Biomechanical strength was measured by L5 compression test using a bone strength tester, TK C Muromachi Kikai Co.
Tokyo, Japan according to our previously published method The rat femur bone was cut from mid-diaphysis with a low-speed diamond blade saw IsoMet; Buehler, Lake Bluff, IL, USA , and after that, the samples were kept in epoxy for nearly 2 h for proper cured and, further samples were polished under the ground Buehler Eco grinder and polisher with the abrasive papers of , , and grit size under the cooling condition and polished with a diamond solution of particle sizes of 1, 0.
After completion of polishing, samples were sonicated for 10 min. The experiment was performed on the T Tribo Indenter Hysitron Inc.
Eight indents with a peak load of µN were applied to cross-section of the bone. The load sequence consists a loading time of 10 s, an unloading segment, and a hold for 10 s.
The resultant load-displacement curve was used to calculate the reduced modulus Er and hardness H by the method of Oliver and Pharr 26 , From the obtained data, we calculated the following parameters: carbonate-to-phosphate ratio area ratio of the carbonate peak [ cm -1 ] to phosphate peak [ cm -1 ] , carbonate-to-amide I ratio area ratio of the carbonate peak [ cm -1 ] to the amide-1 peak [ cm -1 ] and mineral crystallinity ratio intensity ratio of [ to cm -1 ] , which is related to crystal size and stoichiometric perfection.
The amide I band peak contains several sub-peaks that provide information about the collagen matrix and the location of cross-linkage and non-cross linkage. The sub-band of the amide I peak were fited with Gaussian curves at , , , , , and cm -1 using peak analyzing tools OriginPro 8.
Rat cross-linked C-telopeptide of type I collagen CTX-1 kit Cat. E-EL-R and pro-collagen type I N-terminal propeptide PINP kit Cat.
Rat pups 1 to 2-day-old were treated with vehicle or forskolin 1- and 2. After treatment, calvaria were removed and processed for RNA isolation by trizol method qPCR was performed by SYBR green chemistry Thermo Fisher Scientific, Ealtham, MA, USA for the quantitative determination of bone morphogenetic protein 2 BMP2 , type 1 collagen Col I , receptor activator of nuclear factor kappa-B ligand RANKL , and osteoprotegerin OPG as described previously cDNA was synthesized by using 2μg RNA Cat no.
All genes were analyzed using a real-time PCR machine QuantStudio © 3 Real-time PCR Instrument, A , keeping GAPDH as control. Primer sequences are listed below:. Bone marrow was collected from rat femur by flushing out using PBS and cells were measured by a hemocytometer.
After every 48 h media was changed for 21 days. Rat pups 1 to 2-day-old were used to culture calvarial osteoblasts RCO as described previously The adherent cells were treated with CFE 7. RCO were treated with CFE or forskolin for 0 min, 5 min, 15 min, 30 min, 60 min and 90 min.
After treatments, cAMP and cGMP levels were determined by ELISA kits Cayman Co. Data are presented as the mean ± standard error of the mean SEM. One-way ANOVA with a post hoc Tukey test using GraphPad Prism 5 and a significance level of 0.
An unpaired t-test using GraphPad Prism 5 with a significance level of 0. The HPLC method was applied for simultaneous quantification of analytes in CFE.
The chromatograms for the standard mixture and samples are presented in Supplementary Figure 1. The chromatogram of the sample solution obtained in the test for the content of forskolin showed a major peak at a retention time corresponding to that of the forskolin reference standard.
Other diterpene peaks in the sample chromatogram exhibited an additional peak corresponding to isoforskolin. Forskolin and isoforskolin content in the CFE were Osteoblasts are the principal cells that are involved in fracture healing, so we firstly studied the effect of CFE on the differentiation of RCO by ALP activity.
As forskolin is an AC activator, we next studied the effect of CFE Furthermore, at the same concentration, CFE Figure 1 CFE stimulated osteoblast differentiation, cAMP and cGMP in vitro and promoted bone regeneration at the femur osteotomy site.
A RCO were treated with CFE, and differentiation was assessed by ALP assay. B RCO were treated with CFE at the indicated time points and cAMP and C cGMP production were measured. D Adult female rats were treated with vehicle and CFE at indicated doses after femur osteotomy for 12 days, and representative images 10X of calcein deposition at the osteotomy site are shown.
In obese men, mg CFE has been used to study its impact on body mass 5 , 9. Compared with vehicle-treated rats, CFE at all doses significantly increased calcein intensity Figures 1D, E. Trabecular bones at metaphysis and cortical bone parameters were studied using μCT.
N and trabecular thickness Tb. Th compared with control Figure 2B. Cortical parameters including cortical thickness Ct. Th and bone area B.
Ar were significantly increased by CFE over the control Figure 2C. Figure 2 CFE promoted new bone formation in growing rats.
A Femur length. B Representative μCT images left panel and quantitative μCT parameters of the tibia metaphysis right panel. N, trabecular number; Tb. Th, trabecular thickness. C Representative μCT images upper panel and quantitative μCT parameters of the femur and tibia diaphysis lower panel.
Th, Cortical thickness and B. Ar, bone area. D Left panel showing the representative images of double calcein labeling scale bar, µm and histomorphometry parameters right panel of the indicated groups.
E 3-point bending strength of femur was determined by a bone-strength tester. vehicle-treated group. The effect of CFE on bone accrual was measured by dynamic histology by time-spaced calcein labeling study in the periosteal p region of the femur diaphysis. Increase in the surface referent bone formation parameters indicative of increased periosteal apposition complemented our observation of higher Ct.
Th in the CFE group over the control and is likely to afford greater resistance to fracture Accordingly, we measured the bending strength of femur and observed that maximum power and energy to failure were significantly higher in the CFE group compared with control Figure 2E.
The treatment of CFE had no effect on the body weight compared to the vehicle treated group data not shown. Because CFE promoted bone regeneration and stimulated modeling-based bone growth, we speculated that it would have bone conserving effect in OVX model of osteopenia.
At the end of 3 months of treatment, body composition of all groups were assessed by Echo-MRI. Compared with the sham, OVX rats had increased total body mass, lean mass, and fat mass. CFE had no effect on OVX-induced increase in total body mass and lean mass but significantly decreased fat mass Supplementary Figure 2.
We next studied the effects of CFE on appendicular tibia and axial L5 skeletons of OVX rats Figure 3A for representative images. N and Tb. Th were reduced in the OVX group and CFE significantly increased Tb.
Th only. Consequently, trabecular spacing Tb. sp that was increased in the OVX group was significantly reduced by CFE treatment. Th were significantly reversed by CFE with consequent recovery of Tb.
Figure 3B. Figure 3 CFE prevented bone loss in osteopenic rats. A Representative images of tibia metaphysis, and L5 vertebrae are shown.
B Shown are the quantitative μCT parameters of the tibia metaphyses, and L5. Body weight, body mass index BMI , waist and hip circumference, and waist to hip ratio, were monitored fortnightly.
Dietary intake was assessed at the baseline and weeks 4, 8 and Appetite was assessed using visual analogue scales and blood samples were analyzed for plasma lipids, ghrelin, leptin, glucose and insulin at the baseline and end of the intervention.
These findings suggest that C.
Forskolin may promote Injury rehab nutrition guide loss and treat high blood pressure. Tyra Tennyson Francis, MD, is a board-certified dxtract medicine physician and Hydration during breastfeeding serves extrzct the medical director of an Body repair after exercise clinic. Elaine Injury rehab nutrition guide is a registered rxtract, writer, and fact-checker with nearly two decades of experience in educating clients and other healthcare professionals. A natural substance extracted from the Indian coleus plant Coleus forskohliithe supplement forskolin is an herbal remedy related to mint. The plant is found throughout regions of Nepal, Indian, and Southeast Asia. Forskolin is known to activate adenylate cyclase class III AC-IIIan enzyme involved in the regulation of all human cells.
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