Short-term feeding strategies and pork quality

Two experiments were done to determine whether short-term supplementation (5 days pre-slaughter) with magnesium acetate, or a combination of magnesium acetate, tryptophan, vitamin E and vitamin C would improve pork quality. In the first experiment the pigs (Pietrain×Yorkshire, n=96) were fed a standard feed or a magnesium supplemented feed for 5 days prior to slaughter. As a possible stress factor half of the animals were slaughtered upon arrival at the slaughterplant whereas the remaining animals were allowed two hours of rest in lairage before slaughter. Magnesium supplementation did not result in an increase in plasma magnesium concentration at slaughter. Omission of lairage resulted in higher plasma glucose concentrations, but plasma lactate concentrations were not affected. Drip loss and ultimate pH were not affected by diet or omission of lairage. Omission of lairage resulted in poorer color characteristics. This effect was prevented by supplementation with magnesium. In the second experiment the pigs (Pietrain×Yorkshire, n=92) were fed a standard feed or this standard feed supplemented with magnesium acetate, tryptophan, vitamin E and vitamin C for 5 days prior to slaughter. Supplementation with vitamin E did not increase muscle vitamin E concentration. Inclusion of supplements in the diet failed to improve water-holding capacity or color characteristics. These results indicate that short-term supplementation with magnesium acetate, tryptophan, vitamin E and vitamin C is of little value in improving pork quality when pigs are not stressed beyond levels associated with routine slaughter procedures.     

Effect of feeding system on the growth and carcass characteristics of Iberian pigs, and the use of ultrasound to estimate yields of joints

Forty three-castrated male Iberian pigs of the Torbiscal line with an average initial weight of 102.27kg were used. A group of 22 pigs was fed according to the traditional way in free-range conditions with acorn and grass from November to January. Another group of 21 animals was fed 3.72kg/day of a conventional feed that contained 12.53MJME/kg, 16.21% crude protein, 4.94% fat and 6.64% of crude fiber, during the same period. Measurements of backfat thickness and Longissimus dorsi muscle area at the last rib were taken by means of a real-time ultrasound device (Kretz Technick Inc., 600 V-V2.32, Sonovet, Austria) in all pigs before slaughter. After slaughter, carcass, ham and foreleg weights and backfat and muscle area measurement from the last rib chop were recorded. The average daily gain, carcass yield, fat thickness, loin area, ham weight and proportion of hams were significantly higher (P<0.05) in the pigs fed in confinement than in those fed extensively (653.36g, 77.97%, 56.46mm, 3031mm(2), 22.26kg and 14.32% vs 501.48g, 76.94%, 51.74mm, 2673mm(2), 21.52kg and 13.79%, respectively). This paper shows the prediction equations for hams, forelegs and total hams and forelegs weights and proportions with respect to slaughter weight and live ultrasound measurements in pigs fed either extensively or in confinement. The ultrasound measurements better predicted the carcass cuts absolute amounts (R(2)=0.591-0.875) than the carcass cuts proportions (R(2)=0.23-0.40). The use of real-time ultrasound should be used with caution to predict the absolute quantities of hams and forelegs from Iberian pigs since the ultrasound measurements are only moderately accurate.     

The eating quality of liver from pigs fed liquid sugar solution prior to slaughter

Livers from pigs fed 4 lb sugar on the day before slaughter contained more than twenty-five times as much glycogen, 1.8 times as much water and were 1 kg heavier than those from control pigs receiving no carbohydrate (averages of six animals). The pigment concentration was lower resulting in an observable paler raw colour. Livers from sugar fed pigs were more tender and juicy but had a weaker flavour.     

Comparison of different dietary magnesium supplements on pork quality

Forty-eight crossbred (Large White×Landrace) boars were used to compare the effect of dietary magnesium aspartate (MgAsp), magnesium sulphate (MgSO(4)) and magnesium chloride (MgCl(2)) on muscle glycogenolysis and pork quality. The pigs were fed finisher feed supplemented with either MgAsp, MgSO(4) and MgCl(2) for 5 days prior to slaughter. At the abattoir, all pigs received 15 electric shocks from an electric goad 5min prior to slaughter. Pigs fed the diet supplemented with MgSO(4) had the highest plasma Mg concentrations at slaughter in comparison with pigs fed the MgAsp and MgCl(2) supplemented diets. There were no differences in plasma adrenaline and noradrenaline concentrations at slaughter between the different diets. Pigs fed the Mg diets had higher muscle glycogen concentrations in the Longissimus thoracis (LT) muscle at 5min and at 40min (except MgCl(2)) post-slaughter compared to pigs fed the control diet. Also pigs fed the Mg diets had lower muscle lactic acid concentrations in the LT at 5min post-slaughter and lower drip loss at 24hr post-slaughter compared to pigs fed the control diet. These results indicate that cheaper magnesium sources, MgSO(4) and MgCl(2), are as efficacious as MgAsp in reducing drip loss and improving pork quality.     

Manipulation of critical quality indicators and attributes in pork through vitamin E supplementation, muscle glycogen reducing finishing feeding and pre-slaughter stress

The combination of a muscle glycogen reducing diet or a standard diet (control group) with normal (80 mg/kg) and high vitamin E levels (500 mg/kg) and exercise immediately prior to slaughter was used on 56 pigs to investigate the influence on meat quality indicators (pH and temperature) and attributes (drip loss, colour and Warner–Bratzler shear force). The drip loss was reduced in M. longissimus dorsi, M. biceps femoris and M. semimembranosus in pigs given the muscle glycogen reducing diet compared with the control groups, the greatest effect was seen in exercised pigs. These results can be explained by an early post mortem reduction in glycometabolism in pigs fed muscle glycogen reducing diets rather than by an increase in ultimate pH. Noticeably, high dietary vitamin E level increased muscle glycogen stores by about 10% on the day prior to slaughter but not on the day of slaughter in both dietary groups compared with the low dietary vitamin E level, which in fact reduced rather than improved the water-holding capacity, especially in pigs fed the standard diet.     

The significance of diet, slaughter weight and aging time on pork colour and colour stability

The objective of the present study was to investigate the effect of a diet with a low content of digestible starch, slaughter weight and subsequent aging time on meat colour and colour stability. Pork colour was determined as the extent of blooming of M. longissimus thoracis (LT) and M. semimembranosus (SM) after 1, 2, 4, 8 and 15 days postmortem and as colour stability during a subsequent storage period in air for 6 days. Compared to the control diet, the experimental diet resulted in a significantly lower postmortem muscle temperature (1°C; p<0.0001). Moreover, high slaughter weight (110kg) resulted in a higher postmortem temperature in LT (p<0.001) compared to low weight (85kg). Independent of feeding strategy and slaughter weight, the extent of blooming decreased during the first 2-4 days of aging in LT, however, the effect was more pronounced in meat from experimentally fed pigs and pigs with high slaughter weight. This effect was not seen in SM, where a gradual increase in blooming took place throughout the aging period. The colour stability was found to be superior in aged pork from experimentally fed pigs. The discoloration rate was faster in SM compared to LT. In conclusion, the present study shows that the diet composition can be used as a tool to control meat colour and colour stability in pork.     

Effect of exercise and pre-slaughter stress on pork muscle characteristics

The effects of exercise, stress and chill temperature on pork muscle characteristics were studied in a 3 × 2 × 2 factorial experiment in which treatments were assigned to blocks made up of six pigs of the same sex from the same litter. Pigs were fed a corn-soybean meal diet with the two littermate pigs on each treatment fed together. Treatments were (1) a control group receiving no exercise and fed in an 8 m(2) pen; (2) a treatment group fed in an 8 m(2) pen and driven 1·6 km/day for 100 days prior to slaughter; and (3) a treatment group fed in pens with 40 m(2) floor space with feeders placed 5 m from their waterers. When the animals averaged approximately 105 kg in weight, one animal from each pair treated alike was subjected to standardized stress. After slaughter, one side was chilled at 2-3°C and the other at 13-15°C for 24 h. Both sides were then chilled at 2-3°C for an additional 24 h. Exercise did not affect average daily gain (ADG), feed efficiency, yield of total wholesale cuts, muscle pH, protein solubility, fiber diameter and sarcomere length of the Longissimus dorsi (LD) and Quadriceps femoris (QF) muscles or the tenderness of the QF muscle. Exercise decreased backfat thickness and the subjective tenderness of the LD muscle. The effects of stress on the characteristics evaluated were consistent with those that have been previously reported. Chilling temperature and interactions involving chilling temperature did not affect any of the characteristics studied. No exercise × stress interactions were observed. Protein solubility values indicated that pale, soft exudative (PSE) muscle was not a factor in any of the treatments. It was concluded that exercise will produce leaner carcasses but less tender muscle and that exercise will not counteract the effects of pre-slaughter stress.     

Effect of a high linoleic acid diet on lipogenic enzyme activities and on the composition of the lipid fraction of fat and lean tissues in the pig

The aim of this study was to examine the influence of a high linoleic acid diet on lipogenic enzyme activities in pig tissues and on the dietetic quality of these tissues (subcutaneous adipose tissue, Semimembranosus and Diaphragma muscles). Pigs were fed diet M containing 4% of maize oil or diet T containing 4% of tallow from 40 to 100 kg live weight. We did not observe any differences in growth or slaughter performances that could be attributed to diet. Lipogenic enzyme activities were higher in the adipose tissue of pigs fed diet M. However, diet had no effect on lipogenesis in the muscles. The lipid, linoleic acid, and cholesterol contents were higher in the adipose tissue of pigs fed diet M whereas the oleic acid concentration of the adipose tissue was lower. No significant increase in the lipid content of either muscle was however observed in pigs fed diet M. The linoleic acid and cholesterol contents were higher in both muscles of pigs fed diet M than in muscles of pigs fed diet T.     

Transfer of the Meat Aroma Precursors (Dimethyl Sulfide, Dimethyl Disulfide and Dimethyl Trisulfide) from Feed to Cooked Pork

Eight pigs were given standard concentrate feed to a live weight of 70 kg. Subsequently half the pigs were controls and were fed the concentrate feed, while the other half of the pigs were fed an experimental diet consisting of the concentrate feed (89 percent of calories) and Brussels sprouts (11 percent of calories) the last 4 weeks before slaughter. Analyses of volatiles in chops and gastro-intestinal contents were performed using solid-phase micro extraction and subsequent characterization by gas chromatography-mass spectrometry. The ratio between the amount of dimethyl sulfide found in the stomach contents from pigs fed Brussels sprouts and the stomach contents from pigs fed the control diet was 9.0, and correspondingly the ratio was 475.5 for dimethyl disulfide and 8.3 for dimethyl trisulfide. The ratio between the amount of dimethyl sulfide found in meat from pigs fed Brussels sprouts and meat from pigs fed the control diet was 14.9, and correspondingly the ratio was 4.8 for dimethyl disulfide and 1.9 for dimethyl trisulfide. Moreover, several other compounds seem to be influenced by feed composition. Finally, the increase in concentration of sulfur compounds in meat in relation to pork aroma is discussed.     

Study of the branched hydrocarbon fraction of intramuscular lipids from Iberian fresh ham

Branched hydrocarbons from the unsaponifiable lipid fraction of the Biceps femoris muscle of Iberian pigs were analyzed. Fifty-five Iberian pigs were distributed to seven groups according to management system (short Montanera and long Montanera, i.e. fed on acorn and pasture for 55 and 75 days prior to slaughter, respectively, and Pienso, fed on a concentrate diet), and genetic type. A branched hydrocarbon, neophytadiene, was identified only in samples from the pigs fed on the extensive system, with higher levels being found in pigs from the long Montanera group, than in those from the short Montanera group; this compound was not found in the Pienso group. Genotype did not affect the level of branched hydrocarbons found.     

Influence of slaughter weight and stress gene genotype on the water-holding capacity and protein gel characteristics of three porcine muscles

The longissimus lumborum, gluteus medius, and the triceps brachii muscles from 40 animals were used to evaluate the effect of stress gene genotype (non-mutant, NN and mono-mutant, Nn) and live weight at slaughter (110 kg and 140 kg) on the processing quality of fresh pork. The 45 minute and ultimate pH measurements did not differ between genotypes. Total percent protein was not different between samples taken from NN or Nn pigs, nor were there any differences in salt-soluble protein. The M. longissimus lumborum from Nn pigs possessed lower water-holding capacity values and lost greater amounts of water upon cooking. In addition, Nn pigs had lower subjective color and firmness scores which suggest a higher incidence of pale, soft and exudative pork. Slaughter weight did not affect total protein, salt-soluble protein, Minolta L(?), a(?) and b(?) values or subjective color, firmness and marbling scores. Back fat thickness and loineye area increased as slaughter weight increased. Overall, this study suggested that Nn pigs have reduced water retention properties which may result in lower yields in processed meat items. Slaughter weight had limited effects on the processing quality of meat from NN or Nn pigs. There were no interactions of significance between stress gene genotype and slaughter weight, suggesting that the differences in muscle quality and functional properties between NN and Nn pigs are maintained over the slaughter weights used in this study.     

Carry-over of Fusarium toxins (deoxynivalenol and zearalenone) from naturally contaminated wheat to pigs.

The frequent contamination of grain with the Fusarium toxins, deoxynivalenol (DON) and zearalenone (ZON), is an important issue in animal and human nutrition. However, data on the exposure of humans to these toxins through consumption of animal tissues exposed to Fusarium toxins (carry-over) are fragmentary. Therefore, residues of DON, ZON and their metabolites were determined in tissues and body fluids of pigs (female and castrated male) from a fattening trial. Pigs were fed a control (n = 6, 0.24 mg DON and 0.009 mg ZON per kg diet as fed) or a Fusarium toxin-contaminated diet (n = 12, 6.68 mg DON and 0.056 mg ZON per kg diet as fed) either ad libitum or for restrictive consumption for 12 weeks. After slaughter (96.3 +/- 11.6 kg live weight), the concentrations of DON and its metabolite, de-epoxy-DON, were measured in serum, bile, liver, kidney, musculus longissimus and back fat, while ZON and its metabolites, alpha- and beta-zearalenol (alpha-/beta-ZOL), were determined in serum, bile and liver. The mean carry-over factor of DON + de-epoxy-DON, defined as the concentration of both substances in the tissue/fluid divided by the DON concentration in the diet, for all pigs decreased from bile (0.1046 +/- 0.0653) > kidney (0.0151 +/- 0.0070) > liver (0.0057 +/- 0.0043) > serum (0.0023 +/- 0.0018) > muscle (0.0016 +/- 0.0016) > back fat (0.0002 +/- 0.0004). The time interval between the end of feeding and slaughter had no consistent effect on DON + de-epoxy-DON concentrations in the analysed specimen of Fusarium toxin-exposed pigs fed restrictively. No transfer of ZON and its metabolites could be observed into serum of pigs, while the mean carry-over factors of ZON + alpha-ZOL + beta-ZOL were 0.0094 +/- 0.0123 and 4.0 +/- 2.2 for liver and bile, respectively. Therefore, it can be concluded that serum is a reliable indicator for DON exposure, but an inappropriate parameter to deduce ZON exposure, which is better represented by bile concentration of ZON + alpha-ZOL + beta-ZOL. However, the exposure risk to humans by consumption of edible tissues of animals exposed to Fusarium toxins is negligible compared to the direct consumption of grain-based food.     

Strategic finishing feeding as a tool in the control of pork quality

A standard diet and two finishing feeding strategies known to reduce muscle glycogen stores were investigated in combination with exercise immediately prior to slaughter in pigs. The objective was to determine the influence of muscle glycogen at slaughter on temperature and pH in post-mortem muscle, the colour, drip loss and Warner–Bratzler shear force of the meat. The muscle glycogen stores were reduced by strategic finishing feeding. In general, pH_{45 min} was higher in muscles from strategically fed pigs compared with control pigs. Exercise also resulted in higher pH_{45 min} in control pigs compared to non-exercised control pigs, while the opposite was seen in muscles from strategically fed pigs. Exercise resulted in higher muscle temperatures in the carcasses irrespective of feeding strategy. pH_{24 h} were higher in M. biceps femoris and M. semimembranosus from exercised, strategically fed pigs compared with the corresponding controls. In contrast, irrespective of feeding strategy no difference in pH_{24 h} was registered in the meat of non-exercised pigs. Drip loss was lower in meat of strategically fed pigs compared with meat of control pigs. Moreover the drip loss was lowest in the meat of non-exercised pigs. The present study shows that strategic finishing feeding has high potential for the control of pork quality.     

The effect of dietary linoleic acid on the firmness of backfat in pigs of 85 kg live weight

Three groups of 15 female pigs were fed diets containing 0.8%, 1.1%, or 1.8% linoleic acid from 20 to 35 kg followed by 1.0%, 1.2% or 1.4% linoleic acid respectively from 35 kg to slaughter at 85 kg live-weight. Pigs grew at similar rates on all three diets and had similar proportions of carcass fat. The mean proportions of linoleic acid in the backfat lipids were 8.6%, 11.0%, and 13.9% respectively as the amount of the diet increased. There was a significant correlation between the concentration of linoleic acid in the inner backfat lipid and P₂ backfat thickness for those animals on the high and medium linoleic acid diets but not for those on the low diet. Backfat from pigs fed the high linoleic acid diet was softer than that from the other two groups of pigs which could not be distinguished using the subjective finger probe method. Using a mechanical probe backfat consistency was distinguished between all three groups and was inversely correlated (r=?0.775, P<0.001) with the concentration of linoleic acid in the lipid. However, none of the pigs had fat which was too soft for bacon production based on either consistency data or linoleic acid content. If 15% linoleic acid is taken to be the maximum acceptable in bacon, then extrapolation of the regression of linoleic acid concentration on backfat thickness suggested that only the 1.4% linoleic acid diet is likely to result in unacceptably soft fat as pigs become leaner. However, this high concentration, although frequently present in current feeds, is not necessary since it is twice that recommended for normal growth and development of pigs.     

Seroprevalence of anti-hepatitis e virus and anti-salmonella antibodies in pigs at slaughter in Switzerland

Hepatitis E virus (HEV) and Salmonella bacteria are zoonotic pathogens that can be acquired by foodborne transmission because food animals, for example pigs, are recognized as a reservoir. The objectives of this study were to determine the seroprevalence of anti-HEV immunoglobulin G (IgG) and anti-Salmonella antibodies from healthy pigs at slaughter in Switzerland, a country with a good health status of pig herds (e.g., eradication of enzootic pneumonia) compared with those of many countries in the European Union, and a rate of importation of live pigs that is very low (<1%). Based on pooled (diaphragm muscles from 3 to 5 animals per producer) meat juice samples, 120 (60%) of 200 and 8 (4%) of 200 samples were positive for anti-HEV IgG and anti-Salmonella antibodies, respectively. HEV seems to be highly prevalent among fattening pigs in Switzerland, whereas the low seroprevalence of anti-Salmonella IgG has not changed in recent years.     

Dietary Iron in Swine Rations Affects Nonheme Iron and TBARS in Pork Skeletal Muscles

Fifteen crossbred pigs (mean wt = 25 kg) were allocated to three groups and fed to market weight (mean wt = 103 kg) on corn-soy based diets containing either 62,131, or 209 ppm iron. After slaughter, the longissimus dorsi (LD) and rectus femoris (RF) muscles were dissected, cooked, and stored in oxygen-permeable bags for 12 days at 4°C. Cooking increased thiobarbituric acid reactive substances (TBARS) but did not affect nonheme iron (NHI) or α-tocopherol. NHI and TBARS increased continuously during storage while α-tocopherol decreased. NHI and TBARS were higher in cooked pork from pigs fed high-iron diets. Liver iron correlated with muscle iron (p<0.05).     

Prevalence of pigs infected by Salmonella Typhimurium at slaughter after an enterocolitis outbreak

A cross-sectional study was performed to estimate the prevalence of slaughter pigs infected by Salmonella typhimurium after an enterocolitis outbreak in a commercial pig farm, which was characterised by diarrhoea during the growing phase. Anatomopathological and histopathological findings were suggestive of salmonellosis, which was further confirmed by isolation of S. typhimurium from organs and faeces samples from diseased animals. Ileocolic lymph nodes were aseptically collected from 43 pigs during slaughter procedures. The estimated prevalence of Salmonella-infected pigs was 53.48% [confidence interval (CI): 42.94:64.02%]. This finding demonstrates that the carriage of S. typhimurium at slaughter might be high if pigs originate from a batch previously affected by Salmonella-enterocolitis outbreak at the pre-harvest pork production chain.     

Effect of time between adrenaline injection and slaughter on the rate and extent of post-mortem metabolism in porcine skeletal muscle

The aim of this work was to study the effect of time between adrenaline injection and slaughter on the rate and extent of post-mortem metabolism in pig muscle. Five pigs were subcutaneously injected with adrenaline (0·3 mg/kg) or with a saline solution 4 h, 1 h or 15 min prior to slaughter. pH(1), pH(u) and FOP(u) were measured in Longissimus dorsi (LD) and Biceps femoris (BF) muscles. m.LD samples were taken 20 days before slaughter using biopsy sampling, immediately after bleeding and 45 min post mortem for biochemical analysis. m.LD glycolytic potential (very close to glycogen content) was decreased by the injection of adrenaline 4 h and 1 h prior to slaughter, when comparing values at rest (20 days before slaughter) and values determined immediately after bleeding. The depletion was greater fro the injection performed 4 h before slaughter. In this group pH(u) was higher foe adrenaline-injected pigs (5·69 versus 5·47 for pigs injected with saline solution) but the difference was not significant. Pigs injected with adrenaline 1 h prior to slaughter exhibited higher glucose, glucose-6-phosphate and lactate levels immediately after slaughter, lower pH(1) and higher FOP values in m.LD than control pigs. Since the differences in pH(1) were not explained by differences in the rate of build-up of lactate, it was hypothesized that pigs injected with adrenaline 1 h prior to slaughter had lower muscle pH at slaughter. It was concluded that the rate or the extent of post mortem metabolism in pig muscle may be affected independently, by manipulating the time between adrenaline injection and slaughter.     

Dietary β‐cyclodextrin reduces the cholesterol levels in meats and backfat of finishing pigs

BACKGROUND: A hypocholesterolemic effect of β‐cyclodextrin (βCD) has been recently reported in various animals. This study was performed to determine the effect of dietary βCD on hypocholesterolemic effect, and its relationship to bile acid changes of 120 finishing pigs fed a commercial diet containing 0%, 5%, 7%, or 10% βCD for 30 days before slaughter. RESULTS: Plasma total lipids, triacylglyceride, and total cholesterol of βCD supplemented pigs were lower (P < 0.05) than the control pigs. When 5%, 7%, or 10% βCD was supplemented to pigs, cholesterol levels of the plasma were reduced by 26.0%, 28.2%, and 31.3%, respectively. Excretion of total sterol increased (P < 0.05) with increasing βCD supplementation and the composition of excreted bile acid was significantly altered in pigs fed βCD. The cholesterol levels of pig backfat, belly, loin, and ham were reduced as βCD supplementation increased (P < 0.05) in the diet. CONCLUSION: These results suggest that supplementing the finishing diet of swine with βCD resulted in reducing not only circulating lipids and cholesterol but also cholesterol level of pork lean meat and fat, and may provide the method of low‐cholesterol pork production to the swine industry. Copyright © 2008 Society of Chemical Industry     

Electrical stunning by direct brain stimulation in pigs

The effects of electrical stunning of slaughter pigs by electrodes shot into the brain, on sensibility and on the incidence of haemorrhages in the shoulder postmortem, were studied.

To determine the efficiency of this method 37 slaughter pigs were equipped with EEG (electroencephalograph) and ECG (electrocardiograph) electrodes and stunned twice via the EEG electrodes. In the first experiment the pigs were stunned with 25 V and in the second with 25 V followed by a relaxation current. The day after slaughter the shoulders were scored for haemorrhages. Under more practical conditions three groups of 10 pigs each were electrically stunned via electrodes shot into the brain using 25, 50 or 75 V, followed by a relaxation current. The day after stunning the shoulders were scored for haemorrhages.

The pigs did not react to shooting electrodes frontally into the head. After stunning all animals showed a general epileptiform seizure. However, after the second stun the tonic and clonic muscle cramps were impaired. The heart rate increased significantly (P < 0·01) after stunning, and extra systolae and irregular heart rates were observed. In the shoulders of 27% of the carcasses no haemorrhages were found and in the remainder only minor haemorrhages were seen. Under practical conditions a voltage of 87(± 2) V and a current of 511(±20) mA seemed to be necessary for an effective stun. When lower voltages were applied fewer carcasses with shoulder haemorrhages were found.

It may be concluded from these experiments that 25 V (134 mA; 150 Hz) is sufficient for effective stunning of slaughter pigs when electrodes are in direct contact with the brain. But as the optimum position of the electrodes may not be reached under practical conditions a voltage of approximately 90 V (550 mA; 150 Hz) is recommended. After this stunning method only a few haemorrhages were observed in the shoulders.