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This study explored mitochondrial capacities to oxidize carbohydrate and fatty acids and functional optimization of mitochondrial respiratory chain complexes in athletes who regularly train at high exercise intensity (ATH, n = 7) compared with sedentary (SED, n = 7). Peak O2 uptake (V̇o2max) was measured, and muscle biopsies of vastus lateralis were collected. Maximal O2 uptake of saponin-skinned myofibers was evaluated with several metabolic substrates [glutamate-malate (V̇GM), pyruvate (V̇Pyr), palmitoyl carnitine (V̇PC)], and the activity of the mitochondrial respiratory complexes II and IV were assessed using succinate (V̇s) and N,N,N′,N′-tetramethyl-p-phenylenediamine dihydrochloride (V̇TMPD), respectively. V̇o2max was higher in ATH than in SED (57.8 ± 2.2 vs. 31.4 ± 1.3 ml·min−1·kg−1, P < 0.001). V̇GM was higher in ATH than in SED (8.6 ± 0.5 vs. 3.3 ± 0.3 μmol O2·min−1·g dry wt−1, P < 0.001). V̇Pyr was higher in ATH than in SED (8.7 ± 1.0 vs. 5.5 ± 0.2 μmol O2·min−1·g dry wt−1, P < 0.05), whereas V̇PC was not significantly different (5.3 ± 0.9 vs. 4.4 ± 0.5 μmol O2·min−1·g dry wt−1). V̇S was higher in ATH than in SED (11.0 ± 0.6 vs. 6.0 ± 0.3 μmol O2·min−1·g dry wt−1, P < 0.001), as well as V̇TMPD (20.1 ± 1.0 vs. 16.2 ± 3.4 μmol O2·min−1·g dry wt−1, P < 0.05). The ratios V̇S/V̇GM (1.3 ± 0.1 vs. 2.0 ± 0.1, P < 0.001) and V̇TMPD/V̇GM (2.4 ± 1.0 vs. 5.2 ± 1.8, P < 0.01) were lower in ATH than in SED. In conclusion, comparison of ATH vs. SED subjects suggests that regular endurance training at high intensity promotes the enhancement of maximal mitochondrial capacities to oxidize carbohydrate rather than fatty acid and induce specific adaptations of the mitochondrial respiratory chain at the level of complex I.
- metabolism
- exercise training
skeletal muscle is a highly malleable tissue, capable of pronounced metabolic and morphological adaptations in response to contractile activity (i.e., exercise) (17). Endurance training induces marked metabolic and structural adaptations in skeletal muscle, with an increase in mitochondrial mass and capillary density (19). In parallel with these quantitative improvements, some mitochondrial qualitative changes seem to participate in the mitochondrial adaptations, allowing increased exercise performance (40, 45). Our laboratory previously demonstrated that, in addition to increased muscle oxidative capacities, the highest fitness levels are accompanied by a tighter control of respiration by phosphate acceptors and by an increased role of the creatine kinase system, providing a more efficient servo-control of energy production to energy expenditure (43, 45).
Mitochondrial adaptations to endurance training lead to a greater absolute capacity to oxidize all fuel substrates, including carbohydrates (CHO) and fatty acids (FA). The relative contribution of each fuel varies with exercise intensity (4, 8, 30) and training status (5, 9, 20, 21), and CHO are known to be primarily used during high-intensity exercise, independent of fitness level (6, 24, 30). Nevertheless, it has been shown that trained subjects rely less on CHO for fuel than do untrained subjects, even during exercise performed at similar relative high intensity (10, 39). In regard to training adaptations, it is well known that short-term endurance training induces an improvement in the capacity to oxidize lipids during moderate exercise intensity (9, 10, 36). Although endurance training at mixed exercise intensity has been documented to improve mitochondrial function (40), no studies directly compared the relative contribution of each fuel in the mitochondrial function of athletes who trained regularly at a velocity associated with the second ventilatory threshold with sedentary subjects.
Our laboratory recently demonstrated on a murine model the existence of a mitochondrial tissue specificity of substrate oxidation as a function of muscle fiber type (28). Therefore, exploration of the intrinsic mitochondrial capacities to oxidize CHO and lipid may represent a means to visualize the muscular adaptations of metabolic pathways as a function of the training status and training intensity.
Mitochondria oxidize pyruvate and FA into NADH and/or FADH2. These reduced equivalents are oxidized further by the mitochondrial respiratory chain to establish an electrochemical gradient of protons, which is finally used by the ATP synthase to rephosphorylate ADP into ATP. It has been suggested that the increase of some respiratory complex activities could allow the optimization of enzyme ratios along the different mitochondrial respiratory chain complexes (13), which could ameliorate the cellular homeostasis during exercise. However, to date, no study explored the possible adaptations of mitochondrial respiratory chain complexes in human skeletal muscle when muscular activity is highly increased chronically. We hypothesized that, concomitantly with a higher capacity to oxidize CHO, there could be some specific adaptations in the activities of the respiratory complexes in athletes who regularly train at high intensities.
Therefore, this study explored qualitative adaptations at two crucial steps of the mitochondrial machinery in skeletal muscle of athletes (ATH) who trained at high intensity vs. sedentary (SED) human subjects throughout a cross-sectional study. 1) We explored the intrinsic functional capacities of mitochondria to oxidize FA and CHO to determine the long-term adaptations of potentialities for fuel preferences. 2) We investigated the specific adaptations at the level of the mitochondrial respiratory chain complexes.
Seven SED individuals and seven ATH, all men, participated in the study (Table 1). All subjects were informed about the potential risks associated with the experiment before giving their written consent to participate. ATH were engaged in a regular training schedule comprising five training session per week for more than 2 yr, including two weekly training session performed at high intensity. Their respective individual training schedule remained unaltered during the 3 wk preceding the experimental period. The investigation was approved by the Consultant Committee on Human Protection from Biomedical Research of Strasbourg, in accordance with the French Law and with the Declaration of Helsinki. It is important to note that the results are obtained with men, and they cannot be generalized to women.
Study Design
Incremental exercise tests.
Each subject performed an incremental exercise test (IET) to exhaustion to determine the peak power output, peak oxygen uptake (V̇o2max), and ventilatory thresholds. IET were performed in an upright position on an electronically braked cycle ergometer (Medifit 1000S, Belgium). Pedaling frequency was 60–70 revolutions/min and was maintained constant during the test. We used the Hansen equations (15) to determine the maximal theoretical power for SED and ATH subjects and designed the power increments to be individually adjusted so that exhaustion always occur within 12–15 min. The first ventilatory threshold [i.e., lactate threshold (LT)] was determined graphically (2), from a regression analysis of the slope of the carbon dioxide elimination (V̇co2) vs. oxygen consumption (V̇o2) plot. Heart rate and V̇o2 were monitored continuously during the test. Blood lactate was collected immediately at exhaustion and in recovery (1st, 3rd, and 5th min).
Ventilatory parameters.
During the tests, V̇o2 was measured on a breath-by-breath basis using an open-circuit metabolic cart with rapid O2 and CO2 analyses (breath-by-breath metabolic measurement, Sensor Medics, MSE, Yorba Linda, CA). V̇o2max was defined as the highest 30-s average V̇o2 value.
Evaluation of ATH training sessions.
The average training schedule of ATH is given in Table 2 for the 3 wk preceding the study. All ATH were asked to report their individual training schedule into detailed training logs, including duration, distance, and intensity of each training sessions, providing both quantitative and qualitative characterization of the overall training load. Duration and intensity of the training sessions were assessed based on the running velocity spread out in four intensity zones: low exercise intensity < LT < moderate exercise intensity < respiration compensatory point < heavy exercise intensity < V̇o2max < severe exercise intensity.
Metabolic parameters.
During IET tests, 2-ml blood samples were collected into iced tubes for immediate determination of blood LA concentration (Chiron-Diagnostics Serie 800, Bayer, Puteau, France).
Skeletal muscle biopsy.
Vastus lateralis muscle was obtained by the percutaneous Bergström technique after local anesthesia, as previously described (25). Biopsy was taken 48 h after IET, and subjects were asked to refrain from strenuous exercise during this period. The muscle tissue retrieved was rinsed in ice-cold saline and was used for in situ respiration studies.
In situ study of mitochondrial respiration.
The mitochondrial respiration was studied in situ in saponin-skinned fibers. Briefly, fibers were separated under a binocular microscope in solution S at 4°C (see below) and permeabilized in solution S with 50 μg/ml of saponin for 30 min. After being placed 10 min in solution R (see below) to wash out adenine nucleotides and creatine phosphate, skinned separated fibers were transferred in a 3-ml water-jacketed oxygraphic cell (Strathkelvin Instruments, Glasgow, UK) equipped with a Clark electrode, as previously described (21). Solutions R and S contained the following: 2.77 mM CaK2EGTA, 7.23 mM K2EGTA (100 nM free Ca2+), 6.56 mM MgCl2 (1 mM free Mg2+), 20 mM taurine, 0.5 mM DTT, 50 mM potassium-methane sulfonate (160 mM ionic strength), and 20 mM imidazole (pH 7.1). Solution S also contained 5.7 mM Na2ATP, 15 mM creatine-phosphate, while solution R contained 5 mM glutamate, 2 mM malate, 3 mM phosphate, and 2 mg/ml FA free bovine serum. After the experiments, fibers were harvested and dried, and respiration rates were expressed as micromoles of O2 per minute per gram dry weight. Solution R− was similar to solution R without substrates and was used to determined maximal V̇o2 rate for the substrates.
Measurement of the maximal muscular oxidative capacities.
The ADP-stimulated respiration above basal V̇o2 (V̇0) was measured by addition of 2 mM of ADP. After the determination of the V̇0, the maximal fiber respiration rates (V̇max) were measured at 22°C under continuous stirring in the presence of saturating amount of ADP (2 mM) as phosphate acceptor and glutamate-malate as mitochondrial substrates (V̇GM). The acceptor control ratio was V̇GM/V̇0 and represents the degree of coupling between oxidation and phosphorylation.
Measurements of the substrate oxidative capacities.
In separates samples, additions of glycerol-3-phosphate (1.2 mM), pyruvate (1 mM), or palmitoyl-carnitine (135 μM) were done in solution R− in the presence of 2 mM ADP, and maximal V̇o2 rate for these substrates was determined (V̇G3P, V̇Pyr, and V̇PC, respectively). Malate (2 mM) was added with pyruvate and palmitoyl-carnitine to initiate the Krebs cycle.
Measurement of the respiratory chain complexes.
When V̇GM was recorded, electron flow goes through complexes I, III, and IV. Then 4 min after this V̇GM measurement, the complex I was blocked with amytal (2 mM), and then complex II was stimulated with succinate (25 mM). In these conditions, mitochondrial respiration was evaluated by complexes II, III, and IV (V̇S). After that, N,N,N′,N′-tetramethyl-p-phenylenediamine dihydrochloride (TMPD, 0.5 mM) and ascorbate (0.5 mM) were added as an artificial electron donor to cytochrome-c. In these conditions, cytochrome-c oxidase (complex IV) was studied as an isolated step of respiratory chain (V̇TMPD). The ratios V̇S/V̇GM and V̇TMPD/V̇GM allow exploration of complexes I, II, and IV in ATH vs. SED men.
Data are presented as means ± SE. Statistical analyses were performed using Sigma Stat for Windows (version 3.0, SPSS, Chicago, IL). After testing for data distribution normality and variance homogeneity, we used analysis of covariance to adjust group differences for baseline covariates (weight and age). The significance level was set at P < 0.05.
Subject Characteristics
Subjects' characteristics are given in Table 1. As calculated by Hansen's formulas, the predicted V̇o2max was not significantly different between SED and ATH. But by design, the V̇o2max was 89 ± 4% of the predicted V̇o2max, as calculated by Hansen's formula in SED, and 155 ± 4% of predicted V̇o2max in ATH (significantly different from SED, P < 0.001) (41). It is important to notice that, even if ATH trained 79% of total training time at low and moderate intensities, they trained 21% of total training time at very high and severe intensities (Table 2).
Exercise Tests
Table 3 reports the parameters of aerobic performance capacity, as measured during the incremental symptom-limited exercise test. At LT, the V̇o2 expressed in relative values was significantly higher in ATH than in SED (P < 0.001). At peak exercise, V̇o2max was much greater in ATH than in SED, according to the fitness differences, but the respiratory exchange ratio remained similar and >1.1, and LA was >8 mmol/l for both groups, suggesting that all of the subjects reached exhaustion during their IET. As expected, the peak LA was higher in ATH (P < 0.05).
Mitochondrial Substrate Utilization
To compare skeletal muscle oxidative capacities of SED and ATH subjects, we measured the maximal respiratory rates of in situ mitochondria in vastus lateralis samples to determine V̇max with different substrates, as well as the functional activities of the respiratory chain complexes. These parameters allow an exhaustive characterization of muscle maximal oxidative capacities with a given substrate, reflecting both mitochondrial density, as well as the functional properties of skeletal muscle mitochondria.
V̇max with the different mitochondrial substrates are presented in Fig. 1. V̇G3P was low compared with V̇pyr and V̇PC and not significantly different among groups. V̇GM was dramatically higher (+156%) in ATH than in SED subjects (8.6 ± 0.5 vs. 3.3 ± 0.3 μmol O2·min−1·g dry wt−1, P < 0.001). On the other hand, V̇PC was not significantly different between ATH and SED (5.3 ± 0.9 vs. 4.4 ± 0.5 μmol O2·min−1·g dry wt−1, P = 0.37). V̇Pyr was also significantly higher (+58%) in ATH than in SED subjects (8.7 ± 1.0 vs. 5.5 ± 0.2 μmol O2·min−1·g dry wt−1, P < 0.05). When comparing the different substrate pathways, V̇Pyr was significantly higher than V̇PC in ATH (+64%, P < 0.05), which was not the case for SED.
We measured maximal activity of the different complexes of the mitochondrial regulatory chain (Fig. 2). With succinate as electron donor for complex II, V̇S was significantly higher in ATH (+84%) than in SED subjects (11.0 ± 0.6 vs. 6.0 ± 0.3 μmol O2·min−1·g dry wt−1, P < 0.001). Complex IV (TMPD-ascorbate as electron donor) respiratory rates were significantly higher in ATH (+24%) than in SED (20.1 ± 1.0 vs. 16.2 ± 3.4 μmol O2·min−1·g dry wt−1, P < 0.05). The ratios V̇S/V̇GM and V̇TMPD/V̇GM showed the specific adaptations of the mitochondrial complexes I, II, and IV in skeletal muscle of both groups. V̇S/V̇GM (1.3 ± 0.1 vs. 2.0 ± 0.1, P < 0.001), as well as V̇TMPD/V̇GM (2.4 ± 1.0 vs. 5.2 ± 1.8, P < 0.01) were significantly lower in ATH than in SED, suggesting that the excess of complexes II and IV activities compared with complex I activity was proportionally lower in ATH than in SED.
Figure 3 shows the acceptor control ratio (V̇max/V̇0). This parameter was significantly higher in ATH than in SED (5.4 ± 0.5 vs. 3.1 ± 0.5, P < 0.01). These results are in accordance with a previous study (45) and suggest an improvement in the coupling between the electron transport chain and the phosphorylation.
The objective of this study was to compare the mitochondrial function at crucial steps of the mitochondrial machinery between endurance ATH who trained at high exercise intensity and SED subjects. The main results of this study show that the intrinsic mitochondrial capacity to oxidize CHO (i.e., V̇Pyr) is higher in ATH than in SED subjects, whereas the mitochondrial capacity to oxidize lipids (i.e., V̇PC) is not significantly different in both groups. These results support the notion that, besides a quantitative increase in the oxidative capacities, mitochondria also develop adaptations allowing for a specific increase in the mitochondrial capacity to oxidize CHO rather than lipids in ATH who regularly trained at high intensity. Moreover, ATH exhibited lower ratios of complex II and IV to complex I compared with SED subjects, suggesting an optimization of the complex I of the respiratory chain, which could be implicated in the better mitochondrial function observed in ATH.
Regular endurance training induces numerous physiological adaptations, ultimately improving exercise capacity. Previous studies observed alterations in mitochondrial function with endurance training (27, 33, 36, 37, 44, 45). One of the most characteristic adaptations to training is a change in skeletal muscle substrate metabolism (16). Training-induced shift in substrate utilization is classically attributed to the improved muscular oxidative capacities that result from an increase in mitochondrial density (16). However, other factors must also contribute to the training response, as shown in studies in which metabolism (14) or performance was altered (11, 42), despite no change in muscle oxidative capacities. There is a long-standing notion that endurance training causes a shift from predominantly CHO oxidation toward fat oxidation (16). Accordingly, Cogan et al. (9, 10) found that the rate of utilization of plasma glucose is lower in trained than in untrained subjects during moderate- to high-intensity exercise for a given absolute load, and also when the exercise is performed at the same relative (and, therefore, a higher absolute) intensity. For instance, cyclists and triathletes who trained at least 4 times/wk for 2 h or more and had a training history of at least 5 yr show a higher rate of fat oxidation and an increased relative contribution of fat compared with untrained subjects for an exercise at the same relative intensity (60% of V̇o2max) (21). Conversely, O'Brien et al. (26) demonstrate that, during marathon running at 73.3 or 64.5% of V̇o2max, CHO is the primary fuel used. In the same way, Brooks and Mercier (6) have hypothesized that there is a "crossover" effect, such that, during acute exercise at high intensity, the rate of glucose utilization is greater in trained than in untrained subjects, and the rate of glucose uptake increases exponentially with increasing exercise intensity.
In this study, we measured maximal mitochondrial respiration using different substrates, which allow the characterization of mitochondrial fuel preferences for aerobic energy production. Interestingly, we found that mitochondrial intrinsic capacities to oxidize pyruvate (i.e., CHO pathway) are significantly higher than the capacity to oxidize palmitoyl carnitine (i.e., lipid pathway) in ATH who regularly trained at high exercise intensity. These results are in accordance with a recent study of Sahlin et al. (33), which observed that both trained and untrained subject present higher capacity to oxidize pyruvate than palmitoyl carnitine. Moreover, the capacity to use palmitoyl carnitine was not different between ATH and SED. This suggests that mitochondria of ATH exhibited a higher capacity to oxidize CHO than lipid, which was not the case in SED. Indeed, in ATH, maximal mitochondrial respiration with palmitoyl carnitine represents only 60% of the maximal oxidative capacities established with pyruvate and glutamate-malate as substrates. These results substantiate the crossover concept, suggesting that the increased use of CHO during heavy exercise training leads to intrinsic muscular mitochondrial adaptations, promoting the utilization of substrates from the CHO pathways. In fact, high- and very-high-intensity training (20 and 1% of their training duration, respectively) may presumably induce specific metabolic adaptations, different from the adaptations following moderate exercise training. In the same way, the maximal capacity of muscle to transport and phosphorylate glucose during exercise is higher in the trained state, and regular training increases muscle glucose transporter number and hexokinase activity and reduces the intramuscular glucose-6-phosphate concentration during exercise (12). Accordingly, Kjaer et al. (22) reported that the rate of glucose appearance in blood was higher in endurance athletes than in untrained men during exercise at 60–110% of V̇o2max. We found that the functional capacity of mitochondria to oxidize FA was not different between SED and ATH. These results are in line with a molecular study showing similar amounts of mRNAs coding for enzymes involved in the β-oxidation of long-chain FA in tibialis anterior muscle of trained and untrained subjects (34). It is also in accordance with the fact that our ATH, in their competitions, perform at intensities that elicit >80% of V̇o2max, which require mainly CHO oxidation as energy source (6, 38). Thus long-term training at high intensities seems to influence the athlete skeletal muscle profile toward increased mitochondrial capacities to oxidize CHO but not lipids.
It is well known that ATH subjects had a significantly higher myosin heavy chain I percentage and a significantly lower myosin heavy chain IIx percentage than SED subjects (45), and it has been shown that slow-twitch fibers (vs. fast-twitch fibers) have higher activities of enzymes involved in FA oxidation (1). Then we could hypothesize that contractile phenotype could also influence the adaptations of muscular oxidative capacities in our ATH. In case of low-intensity training, substrate oxidation seems to be influenced by fiber-type composition, but not by training status (33). Because our ATH who trained at high intensity increased the capacity to oxidize CHO but not FA, we can suggest that training at high intensity, in contrast to low intensity, promotes muscular CHO oxidation.
Our result could appear in contradiction with the generally well-accepted concept that endurance exercise training leads to a shift of skeletal muscle mitochondria toward an increased use of lipids as a substrate source (for review, see Ref. 18). However, we agree that endurance-trained subjects could use more lipids, both at the same absolute and at the same relative exercise intensity (39), and that higher ADP sensitivity with palmitoyl carnitine than that with pyruvate may influence fuel utilization at low rate of respiration in favor of lipid oxidation (33). Our study provides complementary results, suggesting that mitochondrial adaptations are in favor of higher CHO oxidation capacities in the specific case where subjects regularly train at high intensities. Moreover, further investigations in athletes training at different intensities are needed to explore the role of training intensity in the determination of muscle mitochondrial metabolic profile.
Next to the enhanced capacity to oxidize CHO (i.e., pyruvate), we postulated that high-intensity training could induce some alterations at the level of the mitochondrial respiratory chain complexes to increase the control of mitochondrial respiration (32). The calculation of the ratio of mitochondrial complex activities shows the relative contribution of mitochondrial complexes I, II, and IV in skeletal muscle of both groups. The complex II-to-I ratio, as well as complex IV-to-I ratio were significantly lower in ATH than in SED, suggesting an increase complex I activity compared with complex II and IV in ATH subjects. Because complex I is one of the main limiting steps for the mitochondrial metabolic fluxes in skeletal muscle (32), its augmentation could reflect an improvement in the control coefficient of the respiratory chain (31), allowing amelioration in the rate of NADH oxidation, to support higher CHO oxidation rates. All together, we can postulate that these specific mitochondrial adaptations participate in the increased exercise performance during ATH competition.
Limitations of the Study
Because ATH carried out both moderate and high exercise intensities, we cannot completely rule out that moderate exercise sessions did not participate in the qualitative mitochondrial muscular adaptations. Nevertheless, because of the fact that it has been shown that moderate exercise training preferentially increases fat oxidation capacity (3, 7, 9, 23, 35), together with the fact that muscle glycogen oxidation increased in relation to exercise intensity (30), we can postulate that the muscular adaptations favoring CHO metabolism observed in our ATH were predominantly induced by the high-intensity training sessions. This work is a cross-sectional study rather than a longitudinal study, and thus we cannot definitively state that the high-intensity part of the training program was responsible for the observed alterations. Indeed, there may be a difference in baseline mitochondrial function (before high-intensity training) that would not be captured by this study and could participate in the difference in the mitochondrial function between SED and ATH. Then a longitudinal study exploring the mitochondrial adaptations following high-intensity training needs to be carried out to completely rule out this possibility.
This cross-sectional study suggests that, in addition to higher skeletal muscle oxidative capacities, important qualitative adaptations take place at the level of substrate utilization. In the particular situation where subjects regularly train at high exercise intensity, there seems to be some mitochondrial adaptations favoring the CHO pathway over the lipid pathway for the highest energetic fluxes, and improving the control of mitochondrial metabolic fluxes through improvement of mitochondrial complex I activity compared with complex II and IV.
This research was supported by the Clinical Research Department of Strasbourg's civil hospital and financed by the French Ministry for Heath and Solidarity with a Regional Hospital Protocol of Clinical Research (2002), and by the International Olympic Committee.
↵* F. N. Daussin and J. Zoll contributed equally to this work.
The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
- Copyright © 2008 the American Physiological Society


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Skillet Corn Bread Using Stone Ground Cornmeal
|Vegetable oil
|1 1⁄2 Tablespoon
|All purpose flour
|3⁄4 Cup (12 tbs)
|3 Tablespoon
|Baking powder
|1 Tablespoon
|1 Teaspoon
|1 1⁄4 Cup (20 tbs), preferably stone-ground
|1 Cup (16 tbs)
|2 Large, lightly beaten
|Unsalted butter
|3 Tablespoon, melted
1. Preheat the oven to 425° Warm a 9-inch cast-iron skillet over moderate heat. Add the oil and heat.
2. Meanwhile, in a bowl, sift the flour with the sugar, baking powder and salt. Stir in the cornmeal. Add the milk and eggs and stir lightly. Add the butter and stir just until blended. Scrape the batter into the hot skillet; the oil should bubble. Transfer to the oven and bake for about 18 minutes, or until the center springs back when gently pressed. Turn the corn bread out onto a rack to cool.
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Nutrition Rank
Nutrition Facts
Serving size: Complete recipe
Calories 2047 Calories from Fat 743
% Daily Value*
Total Fat 84 g129.8%
Saturated Fat 34.5 g172.5%
Trans Fat 0 g
Cholesterol 542.3 mg180.8%
Sodium 3429.1 mg142.9%
Total Carbohydrates 287 g95.5%
Dietary Fiber 17.2 g68.7%
Sugars 59.2 g
Protein 46 g92.4%
Vitamin A 37% Vitamin C
Calcium 145.1% Iron 82.6%
*Based on a 2000 Calorie diet


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Ramón Berenguer IV - Lleida
Occupying a strategic position in the shopping and business district of Lleida, opposite the train station, the air-conditioned Ramón Berenguer IV Hotel offers comfortable accommodation in Spain's Catalonia region.
No visit would be complete without admiring the works of art of Lleida, including the magnificent Cathedral de la Seu Vella and the San Martin church. Venture out to explore the Park Segre, an impressive green area which extends along the river.
Highly trained, welcoming, and multilingual staff are happy to help with your vacation needs and requirements, having vast experience in quality hospitality which is offered to visitors from all over the world. The lobby features a television and computer station, and wireless Internet access is complimentary in public areas. Complimentary newspapers are provided in the lobby.
Ramón Berenguer IV - Lleida
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- Luggage storage
- Multilingual staff
- Number of buildings/towers - 1
- Elevator/lift
- Safe-deposit box at front desk
- Library
- Television in common areas
In the room
- Air conditioning
- Hair dryer
- Flat-screen TV
- Cable/satellite TV channels
- Desk
- Free WiFi
- Phone
- In-room safe
Ramón Berenguer IV - Lleida's small print
Also known as
- Ramón Berenguer IV
- Ramón Berenguer IV Hotel
- Ramón Berenguer IV Hotel Lleida
- Ramón Berenguer IV Lleida
Cash transactions at this property cannot exceed EUR 2500, due to national regulations. For further details, please contact the property using information in the booking confirmation.
Mandatory fees
You'll be asked to pay the following charges at check in or check out:
- A tax is imposed by the city: EUR 0.50 per person, per night, up to 7 nights. This tax does not apply to children under 17 years of age.
Optional extras
Self parking costs EUR 7.00 per day
Buffet breakfast is offered for an extra charge of EUR 7.00 per person (approximately)
We have included all charges provided to us by this hotel. However, charges can vary, for example, based on length of stay or the room you book.


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Jack-o'-lantern carved in the shape of Jack from "The Nightmare Before Christmas" and lit with tea lights
PERMISSION TO USE: you are welcome to use this photo free of charge for any purpose including commercial. I am not concerned with how attribution is provided - a link to my flickr page or my name is fine. If the used in a context where attribution is impractical, that's fine too. I want my photography to be shared widely. I like hearing about where my photos have been used so please send me links, screenshots or photos where possible.
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1999 Infiniti I30 Sedan 4dr Sedan
I purchased this car used with 53,000 miles. I now have 75,000 miles after 1 year. It drives very well and its even better on the highway which is what I think it was made for. It has decent pickup as well. It has classy looks for the age of it and for the price you pay for one its much better than a newer domestic car in my opinion. Interior is decent but comfortable. It has been very reliable and I don't for see any problems in the near future. For the bad, The check engine light came on after a month and this is from the Mass air flow sensor. This causes the car to lag in acceleration after starting it up when cold, but if you let it run for a minute it does not cause any problems. After having got rid of it. I can say it is worth a small amount but it is no better than a ford or chevy
We bought this car used in 2004 at 88,000 miles and it now has 174,000 miles 6 years later. This car has been amazing! I nicknamed it the "marshmallow" because it is so comfortable. The maintenance has been quite low $$ wise and reliability amazing! I replaced the original battery last year...10 years later...It still runs so soft and comfortable and performances best at 70 mph. Super comfortable and fun to drive!
heated seats, auto window roll down, sunroof, cushy seats, ski hole (or pass through from trunk to back seats for skis), bose stereo
trunk design is terrible. if it snows or rains the water/snow drops right into your stuff in the trunk. Leather seats get very hot in the sun... you have to remember to put towels or blankets on them.
Bought our I30 as a second family car 3 years ago to get us back and forth from Boston and Florida to visit family. It had 87,000 miles when we bought it and it now has over 150,000. We have spent $0 on maintenance other than the normal oil changes/tires and breaks ect. Liked it so much we went out and bought a G35.
comfort,low cost of ownership, reliability
I agree with the comment on the bose system but for a small price you can change it out for a new model
1999 Infiniti I30 Sedan Touring 4dr Sedan
I bought this car as a my daily driver w/121K miles on it. I put around 15K miles in a year and half. The car drives quiet and smooth. There are all kinds of luxury options that are standard in this car. Everyday I find some new cool feature about this. If you're in the used car market, this is the car. The package you get for the avg. sale price on this car is incredible.
All the luxury options, but esp. BOSE Audio, heated and power seats, smooth ride.
Styling. I guess the G35 took care of that.
This car is as good as they say, no mechanical problems even at 150k. I'm a big guy and the seats do me well. I am comfortable and my back is good even on long trips. This car has it all would buy another in a second.
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PRINT Fantasy Forecaster updated Saturday, Aug. 25 at 9:55 p.m. ET. On tap: The "stretch run" is officially here. Opinions might vary on when that cliched portion of the schedule arrives, but as people tend to measure things based upon the calendar, September, the final "month" of baseball's regular season, seems as good a time as any to reassess your fantasy team's championship prospects. (OK, so there are technically three days of October baseball that are still part of the 2012 regular season, but remember traditional thinking, not to mention statistical splits, measures the concluding month as September.) Sept. 1 arrives on Saturday of the coming week. Barring postponements between now and then, 81 percent of the season will be in the books when play begins that day.
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There are not so many hotel rooms in Toorak, just 3, still everyone can find an offer to suit his choice. The most common offers in this city are hotels, they count 1 out of 3, and their prevalent location is central district. According to our practice, it is the most profitable to book a hotel in the south of Toorak, as it is the location of the hotels with optimal cost and quality offers along with the cheap hotels. Taking into account the small size of the city, there is no need to dwell hotel reservation on the location: the road to the city center would take less then half an hour, so one can book a hotel in any district of Toorak. The most populated with hotels district of Toorak is the south, 55% of cheap hotel rooms are concentrated here. However, even those who want to live in the center can choose from 2 variants of hotel rooms. Toorak is an ideal place for couple and solo, such travellers rate hotels there 7.5 out of 10 on average. Apart from all these, hotels in Toorak offer their customers laundry service, business centre, spa and restaurant/cafe. The most appreciated amenities in Toorak hotels are wifi and service. According to the visitors reviews, hotels in Toorak has fantastic service, great location and good internet service. Take a look at our selection of the cheap hotel rooms in Toorak to make a well-informed choice for your reservation!
Toorak cheap hotels
Guest Houses
All cheap hotels in Toorak
Best hotels in Toorak
Hotels in Toorak are spread around the city within the distance of 2.6 kilometers, while 2 hotels are located directly in the city center. The most popular accommodations in Toorak is
Steam Guest House
located in a 1.2 kilometers from the city center.
The best hotels are here and now. Try them!
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When Veronica Vain walked in on her husband jerkin' it, she never even considered the possibility that her man was just getting a blowjob from his stepdaughter. Caught in the act, Gia Paige hides under the bed while Veronica gets her pussy pounded.
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Aiden Starr might be the worst camp counselor in history, but this busty blonde sure knows how to fuck. When the camp director isn't looking, this curvy babe leans against a tree, and has her pussy fucked while she dirty-talks her way to orgasm town.
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These two girls wanted to be BAD. They broke into professor Mac's house and started pilfering his stuff. Too bad he was filming the whole thing. These little sluts used their tight teen bodies to convince him not to press charges, and fucked him raw!
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Pull the security tapes on the park and you'll see Alina Li getting double-teamed in her car. When she found out these two randos had thick cocks, she stuck her ass out the window so dude could fuck her so raw while she sucked off his friend.
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We heard the park every single night is full of real sluts who want to get pounded by rando cock. So we headed out with a camcorder, and stumbled on this whore Sienna Day. She was so thirsty for fucking, she sucked and fucked down these two guys.
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When you want the job done right, you've got to call a professional. Venice might not exactly be the girl next door, but you'll love watching her put her all-natural body to good use as she sucks and fucks a fat cock all the way to a facial cumshot!
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It's not supposed to be this easy with amateur girls! This skank Chloe was open to anything, willing to show off her sweet-ass titties and even play with herself on camera. Then she let bro fuck her for free right there in her first audition.
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An amateur couple's private session with a Milf sex therapist turned XXX-rated when they realized she was actually porn star Joslyn James. After Sasha learned to deepthroat, this horny pair sent in this tape of their dirty threesome.
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This 18 year old "town whore" got in touch with us cuz she wanted to try making a pro porn scene. We hooked her up with a bro who knows how to bang. Melissa showed off her ace deepthroat skills, and filmed up the fantasy fuck of her nasty d...
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Standing in the bathroom with that juicy booty exposed, Amy teases her boyfriend like a little freak. This horny blonde slut wants that dick in all her holes, and she'll stop at nothing to get that hardcore anal pounding she craves.
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Vocabulary Management SolutionA Vocabulary Management Solution (VMS) provides the tools for a software knowledge management system that can be used on both intranets and extranets for managing taxonomies, thesauri, classification schemes, and provides users with the capability to control files and indexes. Vocabulary Management Solutions will generally provide users with configurable record structures, intranet and extranet client/server networking, Web based user interface for all editorial tasks, and web-based indexing tools that allow for easy searching and browsing.
Stay up to date on the latest developments in Internet terminology with a free weekly newsletter from Webopedia. Join to subscribe now.
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The Fight Nerd had a chance to speak with one of the pioneers of MMA and a man whose never-say-die attitude was most fondly remembered during his days in Pride FC, Gary Goodridge, hwo has just put ou watch video >>
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The Old School MMA Review returns, as we tackle one bizarre early SEG UFC event, UFC 3: The American Dream!Originally billed as champion vs challenger, with the intent to see the returning Ken Shamro watch video >>
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6 things the UFC should add from the Pride Fighting Championships days. watch video >>
The 7 best fighters from Pride Fighting Championships. watch video >>


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Trade Assurance:
Product quality protection
On-time shipment protection
Payment protection
Home > Company Profile
Jingning Qinyuan Fruit And Vegetable Trade Co., Ltd.
Start Order
Onsite Check
The supplier's company premises has been checked by Alibaba.com staff to ensure onsite operations exist there. A third-party verification company has confirmed the legal status of the supplier.
|Business Type:
|Manufacturer, Trading Company
|Gansu, China (Mainland)
|Main Products:
|Fuji Apple,Qinguan Apple,Huaniu Apple,Gala Apple,Fresh Apple
|Total Employees:
|11 - 50 People
|Total Annual Revenue:
|US$1 Million - US$2.5 Million
|Year Established:
|Top 3 Markets:
|Eastern Asia 100.00%
|QUALITY MANAGEMENT SYSTEM CERTIFICATE,The hometown of Chinese apple,Excellent product gold medal,China Famous Fruit,China Green Fruit & Vegetable,China's Top 20 Apple,FSC
|Product Certifications:
Hometown of China Apple Jingning country is located in the west of China, the altitude ...
Hometown of China Apple
Jingning country is located in the west of China, the altitude rough 1344-2245 meters , the sunlight about 2238.2 hours and the annual average temperature 8.3 centigrade.
Our country annual rainfall is 478.3 mm , the features are "Layers deep, four distinct seasons, sufficient sunlight, temperature difference, without pollution".
Jingning Qingyuan Fruit And Vegetable Trade Co., Ltd. was established in 2006. Our company's address is Nanmen village, Renda, Jingning county, Gansu Province, China. Our company covers 6660 square meters, 12000 tons storage in 14 coolstores.
Our main business includes: fruit and vegetable purchase, storage, processing and marketing.
Currently, our orchards rough 27000 square meters, main fruit including Fuji apple, Qinguan apple, Huaniu apple, pears and other fresh fruit. Crisp, delicious with high VC content in our fruit, still now, we have more than 10 countries and district customer s from the world, including Southeast Asia, Middle east, East Europe and so on.
In 2001, the China State Forestry Administration named Jingning as "Hometown of China's apple", and we have won the "China Famous Fruit" title. Jingning Apple has high quality and delicious taste. Welcome to visit our orchards and discuss business cooperation.
Production Capacity View More >
Factory Information
|Product Name
|Units Produced (Previous Year)
|Highest Ever Annual Output
|Unit Type
|Fresh apple
Trade Capacity View More >
Trade & Market
|Main Markets
|Total Revenue(%)
|Eastern Asia
|Export Percentage:
|21% - 30%
|Export Mode:
|Have Own Export License
|No. of Employees in Trade Department:
|1-2 People
Email to this supplier
- Company Overview
- Company Introduction
- Company Capability
- Trade Capacity
- Production Capacity
- Business Performance
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21 November, 2008
a sneak peek...
a sneek peak for next fashion post .. and the clue .. is gonna be this
or this
I still can't decide yet,,
Posted by evita nuh
Friday, November 21, 2008
my passion for fashion
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Doggies Blog: Recent Posts
Maine: Pug Breeders
Below is a sample search of our Pug breeders with puppies for sale. The search tool above returns a list of breeders located listings located nearest to the zip or postal code you enter. From there you can see photos of their pups, find out detailed information on their web page and contact them by email, phonoe or even get driving directions.
(Detailed contact information has been omitted Use Enter your zip or postal code above to access our up to date directory).
This is a FREE search tool! Use the form at the top to get complete listings for breeders in your area!
Small Wonders pugs
Additional Information: PUPPIES are all Sold for 2015!Our black Mercedes has Ch lines and German Ch lines,Fawn stud Cooper is CKC,UKC,and AKC Ch.,with ch lines.no more pups til late 2016 or spring 2017
I am a small home breeder with one breeding AKC female black pug ,black mercedes is 2,puppies are all SOLD,she has ch lines both... ...
Rainbow Pugs
Additional Information: WE WILL NOT HAVE PUPS AVAILABLE TIL LATE 2017, DUE TO A LONG WAIT LIST.
PUPPIES ~ NO PUPS AT THIS TIME. Our pups are delivered in our bedroom, and raised in our living room. Our dogs are family pets, not... ...
KayJo Pugs
Additional Information: I have Two Litters planned for FEBRUARY 2017, Fawns and hopefully some blacks in there. NON-REFUNDABLE DEPOSIT required to hold a pup.6 Deposits in place already.
* CALL OR EMAIL FOR MORE * Please call 207-330-8515 or text, for more info. Go to website: www.kayjopugs.webstarts.com to fill out application... ...
Cold Country Pugs
Additional Information: AKC PUGS,Strong Lines,Shots,Vet Cert,Written Health Guarentee. Many Repeat Customers. Vet References Available Please See Our Website For Further Information. www.coldcountrypugs.com
AKC Reg. dogs. Our pups/dogs all live within our home. I provide a loving, safe enviroment for them. My pups are all born in our... ...
Additional Information: Have a litter due in middle of September.
Additional Information: I have a litter of pug puppies available for deposits now have been breeding pugs for over 20 years I have 3 fawn males 1 fawn female 1 black female vet check AKC papers born 4/18 ready June 13th
PLEASE READ AD CAREFULLY BEFORE EMAILING No puppies available until 2017 My wait list is now pretty full, I will add people for next year... ...
Gamekeeper Kennel
Additional Information: 50 years of breeding we assure satisfaction.Visits are encouraged and welcome by appointment.Pups are suitable for pet,show or breeding,no congenital issues.Phone calls preferred.
Pug Puppy Breeder of Boston
Additional Information: Champion bloodlines, litters expected Summer 2016. Fawn & Black healthy, lovable pug puppies. Email phone # to be added to my wait list. Many references on my website. https://pugpuppy.shutterfly.com
Welcome to Pug Puppy Breeder of Boston. Home of quality, well cared for AKC Registered - Champion Lined Pugs. Our Pug puppies and dogs... ...
Additional Information: NO PUPPIES AVAILABLE 2/13/16.ALL OUR PUGS WERE RECENTLY TESTED FOR P.D.E. AND ARE ALL NON-CARRIERS.
We breed Pugs for temperament and quality. We try to breed the closest we can to the AKC regulations. Our puppies are from AKC champion blood... ...
Additional Information: AKC Pug Puppies.My pugs are all Champion Sired & paired to produce the nicest puppies to the pug standard.I breed Blacks & Fawns for health & quality with fabulous pedigrees.Puppies due April 2015
Healthy Pug pups bred for show or companion.Raised in home from great AKC Champion lines as well as best care & food to provide... ...
Murphy's Pugs
Additional Information: no puppies at this time litter is sold
We are a small breeder of pugs located in Warren, Ma. We have been breeding pugs for 26 years now with much pride and satisfaction. � We usually... ...
Gifford's Pugs
Additional Information: Beautiful Male puppy available DOB 7-23-2016....
Here at the Giffords all our adults and pups are raised in the house with access to a fully fenced in yard. No kennels..... All our pugs are part... ...
Additional Information: 🐾 sorry no pups Available at this time Please check back in a month
Hello there, I live on a mini farm with chickens, Horses, ponies ,1 cat right now and of course Pugs. I have been breeding AKC REG>pugs for... ...
Additional Information: Sorry, no pups available at this time and none on the way. Will post if and when we have puppies next!
Pepper's Perfect Pugs
Additional Information: Please contact me for information, email is best, all pups include dew claws removed, first shots, no papers, flea/tick treatment, vet check and puppy pack...........$575
No pups at the moment, litter due mid October. My pups do not come with papers, that is why I can offer them at a reasonable price... ...
Andi Pugs
Additional Information: We do not have any puppies available at this time.We breed top-quality AKC show pugs. Our dogs have extensive health testing done, and most are conformation Champions.
Cindy's Pugs
Additional Information: All our pups have found their furever homes. We won't be expecting again until spring 2017. Please feel free to check back with us then or e mail me if you have any questions.
We have been breeding these adorable pugs for over 10 years. All puppies are raised under foot with lots of love and we have both... ...
Additional Information: SORRY NO BABIES AVAILABLE AT THIS TIME
joan moore
Additional Information: We do not have any pups at this time, please watch for updates!
Our pups are home raised and loved. Will be started on wee wee pads. Our pups come with AKC pet pedigree papers, one year health guarantee and... ...
Country Girlz Pugz
Additional Information: Family owned, dedicated AKC hobby pug breeder. All our dogs and pups are raised in our home not a kennel.Our dogs are all pre-spoiled, socialized up to date on shots,vet checked.
We our proud to announce, that our black female Darla,and Brindle male Ranger, have become parents of six gorgeous puppies on... ...
Christina's Angel's/Panini Pugs
Additional Information: We were happy to welcome Ruckus and Storms litter on 9/19/16. There are only two platinum puppies still available, so call today for Your little puglet!! Check out our facebook page for current pics
We are a private breeder here in Northern N.J, producing standard and rare pugs of color. we produce Black, Fawn, Apricot (reddish fawn),... ...
Additional Information: AKC PUG PUPS ALL COLORS, FAWN, BLACK, APRICOT, SILVER/GRAY, WHITE, BRINDLES. OUR PUGS ARE ADORABLE AND ARE USED FOR ADDS AND COMMERCIALS ALSO OFFER A 5YR HEALTH GUARANTEE
Additional Information: I have 1 reg. black female $1,800 2 reg. black males $1,700 1 reg. fawn male $1,700. text or e-mail 484-796-1064.1 Pug/Eng/French brindle bull dog mix.$900.
Casull Pugs
Additional Information: **AKC reg'd Fawn puppies. **AKC Champion males available for stud service. **Retired adult Pugs available sometimes. **Also some rescue Pugs who need homes at times.
Puppies range from $1,500 - 2,500. There is an application process and a spay/neuter contract. Fawn litter born 4/3/2016 are Champion... ...
Additional Information: PUPPIES DUE MID JUNE 2016BREEDERS OF EXCEPTIONAL QUALITY AKC REGISTERED PUGS FOR OVER 30 YEARS. SOME OF THE TOP WINNING PUGS OF ALL TIME ARE IN OUR PEDIGREe
Super Cooper
Additional Information: I only provide stud service. Cooper is a great stud! He is a very friendly, fun loving dog with an awesome temperament. His pups are on the small side averaging 18 lbs. $500 is his fee.
Cooper is a happy, healthy, absolutely adorable 1 1/2 year old male pug with awesome markings and great genes! He is energetic,... ...
Happy Pugs
Additional Information: All puppies come with first shots and dewormed. They are health guaranteed and raised with children, dogs and cats. We do NOT ship. It is too high of a risk.
Last little guy left waiting to go home! We only have one brindle male pug available! He is $1400. They are home raised with kids, other dogs,... ...
Dogtown Pugs
Additional Information: Welcome to the site of the Dogtown Pugs! We are located in the rural mountainous area of Northeast Pennsylvania--but don't let the word "rural" frighten you! We are not far from the interestates I-80 and I-81. We have raised many animals (chickens, ducks, geese, cats, rescued dogs, rabbits, honey bees...etc) but in 2004, our lives changed when three pugs were made part of our family. We've heard many snorts, snores and sneezes but we've grown to love their noises and humorous, yet sweet personalities. We hope you can find a pug to add humor and a new lasting friendship to your life, along with a lot of love and kisses.
We are located in the rural mountainous area of Northeast Pennsylvania--but don't let the word "rural" frighten you! We... ...
Wildflower's Pugs
Additional Information: text 484-796-1064 I have 1 reg. Blk. Female $1,800.2 reg. Blk. males,1 fawn male $1,700Come w/health cert. de-wormed,shots.and and a little spoiled. 1 Pug/ fr/eng. bull dog mix.$900.Brindle
Important: breeders.NET has not screened the listed breeders and accepts no responsibility for their reputation or quality. It is your responsibility to investigate. See How to Find a Dog.
New Articles
HEALTH Summer is coming on fast, so it's time to plan how you will keep your dog safe and healthy through the lazy, carefree, warm days.
NUTRITION With the wide variety of food at Thanksgiving dinner, chances are you'll want to give your dog something special, too. If you're contemplating what to feed your dog for the holiday, here is a guide to a great Canine Thanksgiving Feast.
DOG HEALTH So you have your new puppy picked out. There are quite a few shots, treatments and examinations that will keep the newest member of your family healthy.
DOG FUN Walking your dog is not only crucial to keeping him healthy and happy, it strengthens the bond between your canine friend and his caregiver. There are a lot of obstacles out there. Don't forget these simple tips to keep your walk fun and safe in the outside world.
HEALTH The same techniques that physiotherapists use to treat a variety of injuries and conditions in humans have been adapted to suit animals with great success. Family pets, show dogs, and working dogs can all benefit greatly from physiotherapy. Dogs whose activities involve a lot of agility are especially susceptible to the types of problems that physiotherapy can address.
FIRST TIME OWNERSBringing a dog into your family is a decision where many people don't realize it's magnitude until after they have the dog. There are a number of things that you need to research before you decide to purchase a dog, and it starts right in your own home.
HEALTH Many believe that a dog and a new baby cannot happily coexist, so therefore the dog has to go. This is not necessarily the case. A new baby does not mean you have to abandon your dog.
DOG TRAINING It's not enough to simply walk the dog through the front door and hope he will acclimate to his new surroundings; you have to teach him how to interact with the new people and items in his environment.


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Economic development is a process of continuous technological innovation and structural transformation. Development thinking is inherently tied to the quest for sustainable growth strategies. This book provides a neoclassical approach for studying the determinants of economic structure and its transformation and draws new insights for development policy. The market is the basic mechanism for effective resource allocation at each level of development. However, economic development as a dynamic process entails structural changes, including industrial upgrading and diversification and corresponding improvements in hard and soft infrastructure. Such upgrading and improvements require coordination and go hand in hand with large externalities to firms' transaction costs and returns to capital investment. Thus, in addition to an effective market mechanism, the government should play an active role in facilitating structural changes. The book provides empirical evidence in support of this framework as well as concrete advice to development practitioners.
You can read this item using any of the following Kobo apps and devices:


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The simplest, most effective way to create a collaborative school self-evaluation and strategic action plan
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Empower your Middle Leaders with a process to help them evaluate their departments and create action plans for improvement
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This has been repeated to death on smashboards.com
When brawl comes out, I assure you the better place to get people's contact info etc would be on smashboards. But the people who join on smashboards tend not to play with items, have 4 stock battles and are very skilled in melee.
Sign me up and we'll both be surprised as to how well we all are in Brawl when it comes out.