The effect of a large freshwater perturbation on the glacial north Atlantic Ocean using a coupled general circulation model

The commonly held view of the conditions in the North Atlantic at the last glacial maximum, based on the interpretation of proxy records, is of large-scale cooling compared to today, limited deep convection, and extensive sea ice, all associated with a southward displaced and weakened overturning thermohaline circulation (THC) in the North Atlantic. Not all studies support that view; in particular, the "strength of the overturning circulation" is contentious and is a quantity that is difficult to determine even for the present day. Quasi-equilibrium simulations with coupled climate models forced by glacial boundary conditions have produced differing results, as have inferences made from proxy records. Most studies suggest the weaker circulation, some suggest little or no change, and a few suggest a stronger circulation. Here results are presented from a three-dimensional climate model, the Hadley Centre Coupled Model version 3 (HadCM3), of the coupled atmosphere - ocean - sea ice system suggesting, in a qualitative sense, that these diverging views could all have occurred at different times during the last glacial period, with different modes existing at different times. One mode might have been characterized by an active THC associated with moderate temperatures in the North Atlantic and a modest expanse of sea ice. The other mode, perhaps forced by large inputs of meltwater from the continental ice sheets into the northern North Atlantic, might have been characterized by a sluggish THC associated with very cold conditions around the North Atlantic and a large areal cover of sea ice. The authors' model simulation of such a mode, forced by a large input of freshwater, bears several of the characteristics of the Climate: Long-range Investigation, Mapping, and Prediction (CLIMAP) Project's reconstruction of glacial sea surface temperature and sea ice extent.

Empirical evidence for a nonlinear effect of galactic cosmic rays on clouds

Galactic cosmic ray (GCR) changes have been suggested to affect weather and climate, and new evidence is presented here directly linking GCRs with clouds. Clouds increase the diffuse solar radiation, measured continuously at UK surface meteorological sites since 1947. The ratio of diffuse to total solar radiation-the diffuse fraction, (DF)-is used to infer cloud, and is compared with the daily mean neutron count rate measured at Climax; Colorado from 1951-2000, which provides a globally representative indicator of cosmic rays. Across the UK, oil days of high cosmic ray flux (above 3600 X 10(2) neutron counts h(-1), which occur 87% of the time on average) compared with low cosmic ray flux, (i) the chance of an overcast day increases by (19 +/- 4)%; and (ii) the diffuse fraction increases by (2 +/- 0.3)%. During sudden transient reductions in cosmic rays (e.g. Forbush events), simultaneous decreases occur in the diffuse fraction. The diffuse radiation changes are; therefore; unambiguously due to cosmic rays. Although the statistically significant nonlinear cosmic ray effect is small, it will have a considerably larger aggregate effect on longer timescale (e.g. centennial) climate variations when day-to-day variability averages out.

Effect of dietary supplementation of different oils during the first or second half of pregnancy on the glucose tolerance of the sow

Poor glucose tolerance may be an under-researched contributory factor in the high (10% to 20%) pre-weaning mortality rate observed in pigs. Insulin resistance commences at around week 12 of gestation in the sow, although there are conflicting reports in the literature about the extent to which insulin resistance is modulated by maternal diet. The aim of the study was to determine the effects of supplementing the maternal diet with different dietary oils during either the first half or the second half of gestation on the glucose tolerance of the sow. Sows were offered the control (C: n = 5) diet as pellets or the C diet plus 10% extra energy (h = 16 per group) derived from either. (i) extra pellets; (ii) palm oil; (iii) olive oil; (iv) sunflower oil; or (v) fish oil. Experimental diets were fed during either the first (G1) or second (G2) half of gestation. A glucose tolerance test (GTT) was conducted on day 108 of gestation by administering 0.5g/kg glucose i.v. Blood samples were taken every 5 to 10 min for 90 min post administration. The change in body weight and backfat thickness during gestation was similar but both type and timing of dietary supplementation influenced litter size and weight. With the exception of the sunflower oil group, supplementing the maternal diet in G1 resulted in larger and heavier litters, particularly in mothers offered palm oil. Basal blood glucose concentrations tended to be more elevated in G1 than G2 groups, whilst plasma insulin concentrations were similar Following a GTT, the adjusted area under the curve was greater in G1 compared to G2 sows, despite no differences in glucose clearance. Maternal diet appeared to influence the relationship between glucose curve characteristics following a GTT and litter outcome. In conclusion, the degree of insulin sensitivity can be altered by both the period during which maternal nutritional supplementation is offered and the fatty acid profile of the diet.

Effect of replacing calcium salts of palm oil distillate with rapeseed oil, milled or whole rapeseeds on milk fatty-acid composition in cows fed maize silage-based diets

Inclusion of rapeseed feeds in dairy cow diets has the potential to reduce milk fat saturated fatty acid (SFA) and increase cis-monounsaturated fatty acid (cis-MUFA) content but effectiveness may depend on the form in which the rapeseed is presented. Four mid-lactation Holstein dairy cows were allocated to four maize silage-based dietary treatments according to a 4 x 4 Latin Square design, with 28-day experimental periods. Treatments consisted of a control diet (C containing 49 g/kg dry matter (DM) of calcium salts of palm oil distillate (CPO), or 49 g/kg DM of oil supplied as whole rapeseeds (WR), rapeseeds milled with wheat (MR) or rapeseed oil (RO). Replacing CPO with rapeseed feeds had no effect (P > 0.05) on milk fat and protein content, while milk yields were higher (P < 0.05) for RO and MR compared with WR (37.1, 38.1 and 34.3 kg/day, respectively). Substituting CPO with RO or MR reduced (P < 0.05) milk fat total SFA content (69.6, 55.6, 71.7 and 61.5 g/100g fatty acids for C, RO, WR and MR, respectively) and enhanced (P < 0.05) milk cis-9 18:1 MUFA concentrations (corresponding values 18.6, 24.3, 17.0 and 23.0 g/100g fatty acids) compared with C and WR. Treatments RO and MR also increased (P < 0.05) milk trans-MUFA content (4.4, 6.8, 10.5 g/100g fatty acids, C MR and RO, respectively). A lack of significant changes in milk fat composition when replacing CPO with WR suggests limited bioavailability of fatty acids in intact rapeseeds. In conclusion, replacing a commercial palm oil-based fat supplement in the diet with milled rapeseeds or rapeseed oil represented an effective strategy to alter milk fatty acid composition with the potential to improve human health. Inclusion of processed rapeseeds offered a good compromise for reducing milk SFA and increasing cis-MUFA, whilst minimising milk trans-MUFA and negative effects on animal performance.

Effect of dietary vitamin A restriction on marbling and conjugated-linoleic acid (CLA) content in Holstein steers

To determine the effect of duration of dietary vitamin A restriction on site of fat deposition in growing cattle, 60 Holstein steers (BW = 218.4 ± 6.55 kg) were fed a diet based on high-moisture corn with 2,200 IU supplemental vitamin A/kg DM (C) or no supplemental vitamin A for a long (243 d; LR) or short (131 d; SR) restriction prior to harvest at 243 d. The SR steers were fed the C diet for the first 112 d. Steers were penned individually and fed for ad libitum intake. Jugular vein blood samples for serum retinol analysis were collected on d 1, 112, and 243. Carcass samples were collected for composition analysis. Subcutaneous fat samples were collected for fatty acid composition. Fat samples from the i.m. and s.c. depot were collected to measure adipocyte size and density. Feedlot performance (ADG, DMI, and G:F) was not affected (P > 0.05) by vitamin A restriction. On d 243, the i.m. fat content of the LM was 33% greater (P < 0.05) for LR than for SR and C steers (5.6 vs. 3.9 and 4.2% ether extract, respectively). Depth of back fat and KPH percentage were not affected (P = 0.44 and 0.80, respectively) by vitamin A restriction. Carcass weight, composition of edible carcass, and yield grade were similar among treatments (P > 0.10). Liver retinol (LR = 6.1, SR = 6.5, and C = 44.7 µg/g; P < 0.01) was reduced in LR and SR vs. C steers. On d 243, LR and SR steers had similar serum retinol concentrations, and these were lower (P < 0.01) than those of C steers (LR = 21.2, SR = 25.2, and C = 36.9 µg/dL). Intramuscular adipose cellularity (adipocyte/mm2 and mean adipocyte diameter) on d 112 and d 243 was not affected (P > 0.10) by vitamin A restriction. Restricting vitamin A intake for 243 d increased i.m fat percentage without affecting s.c. or visceral fat deposition, feedlot performance, or carcass weight. Restricting vitamin A intake for 131 d at the end of the finishing period appears to be insufficient to affect the site of fat deposition in Holstein steers.

Pea-barley intercropping and short-term subsequent crop effects across European organic cropping conditions

Grain legumes are known to increase the soil mineral nitrogen (N) content, reduce the infection pressure of soil borne pathogens, and hence enhance subsequent cereals yields. Replicated field experiments were performed throughout W. Europe (Denmark, United Kingdom, France, Germany and Italy) to asses the effect of intercropping pea and barley on the N supply to subsequent wheat in organic cropping systems. Pea and barley were grown either as sole crops at the recommended plant density (P100 and B100, respectively) or in replacement (P50B50) or additive (P100B50) intercropping designs. In the replacement design the total relative plant density is kept constant, while the additive design uses the optimal sole crop density for pea supplementing with 'extra' barley plants. The pea and barley crops were followed by winter wheat with and without N application. Additional experiments in Denmark and the United Kingdom included subsequent spring wheat with grass-clover as catch crops. The experiment was repeated over the three cropping seasons of 2003, 2004 and 2005. Irrespective of sites and intercrop design pea-barley intercropping improved the plant resource utilization (water, light, nutrients) to grain N yield with 25-30% using the Land Equivalent ratio. In terms of absolute quantities, sole cropped pea accumulated more N in the grains as compared to the additive design followed by the replacement design and then sole cropped barley. The post harvest soil mineral N content was unaffected by the preceding crops. Under the following winter wheat, the lowest mineral N content was generally found in early spring. Variation in soil mineral N content under the winter wheat between sites and seasons indicated a greater influence of regional climatic conditions and long-term cropping history than annual preceding crop and residue quality. Just as with the soil mineral N, the subsequent crop response to preceding crop was negligible. Soil N balances showed general negative values in the 2-year period, indicating depletion of N independent of preceding crop and cropping strategy. It is recommended to develop more rotational approaches to determine subsequent crop effects in organic cropping systems, since preceding crop effects, especially when including legumes, can occur over several years of cropping.

Effect of energy and protein supplementation on phosphorus utilization in lactating dairy cows

Two experiments were undertaken in which grass silage was used in conjunction with a series of different concentrate types designed to examine the effect of carbohydrate source, protein level and degradability on total dietary phosphorus (P) utilization with emphasis on P pollution. Twelve Holstein-Friesian dairy cows in early to mid-lactation were used in an incomplete changeover design with four periods consisting of 4 weeks each. Phosphorus intake ranged from 54 to 80 g/day and faecal P represented the principal route by which ingested P was disposed of by cows, with insignificant amounts being voided in urine. A positive linear relationship between faecal P and P intake was established. In Experiment 1, P utilization was affected by dietary carbohydrate type, with an associated output of 3.3 g faecal P/g milk P produced for all treatments except those utilizing low degradable starch and low protein supplements, where a mean value of 2.8 g faecal P/g milk P was observed. In Experiment 2, where two protein levels and three protein degradabilities were examined, the efficiency of P utilization for milk P production was not affected by either level or degradability of crude protein (CP) but a significant reduction in faecal P excretion due to lower protein and P intake was observed. In general, P utilization in Experiment 2 was substantially improved compared to the Experiment 1, with an associated output of 1.8 g faecal P/g milk P produced. The improved utilization of P in Experiment 2 could be due to lower P content of the diets offered and higher dry matter (DM) intake. For dairy cows weighing 600 kg, consuming 17-18 kg DM/day and producing about 25 kg milk, P excretion in faeces and hence P pollution to the environment might be minimized without compromising lactational performance by formulating diets to supply about 68 g P/day, which is close to recent published recommended requirements for P.

Effect of replacing calcium salts of palm oil distillate with extruded linseeds on milk fatty acid composition in Jersey and Holstein cows

Clinical and biomedical studies have provided evidence for the critical role of n-3 fatty acids on the reduction of chronic disease risk in humans, including cardiovascular disease. In the current experiment, the potential to enhance milk n-3 content in two breeds with inherent genetic differences in mammary lipogenesis and de novo fatty acid synthesis was examined using extruded linseeds. Six lactating cows (three Holstein and three Jersey) were used in a two-treatment switchback design with 3 × 21-day experimental periods to evaluate the effect of iso-energetic replacement of calcium salts of palm oil distillate (CPO) in the diet (34 g/kg dry matter (DM)) with 100 g/kg DM extruded linseeds (LIN). For both breeds, replacing CPO with LIN had no effect (P > 0.05) on DM intake or milk yield, but reduced (P < 0.05) milk fat and protein yield (on average, from 760 to 706 and 573 to 552 g/day, respectively). Relative to CPO, the LIN treatment reduced (P < 0.01) total saturated fatty acid content and enhanced (P < 0.001) 18:3n-3 in milk, whereas breed by diet interactions were significant for milk fat 16:0, total trans fatty acid and conjugated linoleic acid concentrations. Increases in 18:3n-3 intake derived from LIN in the diet were transferred into milk with a mean marginal transfer efficiency of 1.8%. Proportionate changes in milk fatty acid composition were greater in the Jersey, highlighting the importance of diet–genotype interactions on mammary lipogenesis. More extensive studies are required to determine the role of genotype on milk fat composition responses to oilseeds in the diet.

Effect of replacing grass silage with maize silage in the diet on bovine milk fatty acid composition

Even though extensive research has examined the role of nutrition on milk fat composition, there is less information on the impact of forages on milk fatty acid (FA) composition. In the current study, the effect of replacing grass silage (GS) with maize silage (MS) as part of a total mixed ration on animal performance and milk FA composition was examined using eight multiparous mid-lactation cows in a replicated 4 X 4 Latin square with 28-day experimental periods. Four treatments comprised the stepwise replacement of GS with MS (0, 160, 334 and 500 g/kg dry matter (DM)) in diets containing a 54:46 forage: concentrate ratio on a DM basis. Replacing GS with MS increased (P < 0.001) the DM intake, milk yield and milk protein content. Incremental replacement of GS with MS in the diet enhanced linearly (P < 0.001) the proportions of 6:0-14:0, decreased (P < 0.01) the 16:0 concentrations, but had no effect on the total milk fat saturated fatty acid content. Inclusion of MS altered the distribution of trans-18:1 isomers and enhanced (P < 0.05) total trans monounsaturated fatty acid and total conjugated linoleic acid content. Milk total n-3 polyunsaturated fatty acid (PUFA) content decreased with higher amounts of MS in the diet and n-6 PUFA concentration increased, leading to an elevated n-6: n-3 PUFA ratio. Despite some beneficial changes associated with the replacement of GS with MS, the overall effects on milk FA composition would not be expected to substantially improve long-term human health. However the role of forages on milk fat composition must also be balanced against the increases in total milk and protein yield on diets containing higher proportions of MS.

Effect of acute administration of recombinant human leptin during the neonatal period on body temperature and endocrine profile of the piglet

Background: Leptin is produced predominantly by white adipocytes; in adults it regulates appetite and energy expenditure but its role in the neonate remains to be fully established. Objectives: To examine the effects of acute administration of recombinant human leptin on the endocrine profile and thermoregulation of neonatal pigs. Methods: 24 pairs of siblings (n = 48) were administered with either a single dose (4 mu g ml(-1) kg(-1) body weight) of leptin (L: n = 24) or a placebo (P: n = 24) on day 6 of neonatal life. Rectal temperature was recorded, and tissue samples were taken at 1 (n = 12), 2 (n = 12), 4 (n = 12) or 6 (n = 12) hours post-administration. Plasma concentrations of hormones and metabolites were determined in conjunction with messenger RNA (mRNA) for leptin and uncoupling protein-2. Results: Plasma leptin increased following leptin administration, and differences in concentrations of insulin, thyroxine and non-esterified fatty acids were observed between the two groups. Initially, rectal temperature decreased in L pigs but returned to start values by 1.5 h. This decline in rectal temperature was delayed in placebo animals, resulting in differences between treatments at 1.5 and 2 h. Conclusions: Acute leptin administration alters the endocrine profile of pigs and influences the thermoregulatory ability of the neonate. Copyright (C) 2007 S. Karger AG, Basel.

Environmental control for layers

Light patterns have less effect on numbers of eggs laid by current stocks than on those of forty years ago, but the principles have not changed. Ovarian activity is stimulated by increasing photoperiods and suppressed by decreasing photoperiods. The light pattern used during rearing can still have large effects on age at 50% lay, even for modern stocks. Early sexual maturity maximises egg numbers but gives smaller eggs. Late maturity maximises egg size at the expense of numbers. The relationship between egg output (g/hen d) and age at first egg is curvilinear, with maximum yield occurring in flocks maturing in about the centre of their potential range. Fancy patterns of increasing daylength after maturity are probably not justified. A flock held on a constant 14h day will lay as many eggs as one given step up lighting. Intermittent lighting saves about 5% of feed consumption with no loss of output, provided that the feed has adequate amino acid content to allow for the reduced feed intake. Producers with light-proof laying houses should be taking advantage of intermittent lighting. The recommended light intensity for laying houses is still 10 lx, although the physiological threshold for response to changes in photoperiod is closer to 2 lx. Very dim (0.05 lx) light filtering into blacked out houses will not stimulate the hypothalamic receptors responsible for photo-sexual responses, but may affect the bird's biological clock, which can alter its response to a constant short photoperiod. Feed intake shows a curvilinear dependence on environmental temperature. At temperatures below the panting threshold, performance can be maintained by adjusting the feed so as to maintain an adequate intake of critical amino acids. Above the panting threshold, the hen is unable to take in enough energy to maintain normal output. There is no dietary modification which can effectively offset this problem. Diurnally cycling temperatures result in feed intake and egg production equivalent to that observed under a constant temperature equal to the mean of the cycle. When the poultry house is cooler at night than by day, it helps to provide light so that the birds can feed during the cooler part of the cycle.

Short Communication: effects of 2-hydroxy-4-(methylthio) butanoic acid isopropyl ester on milk production and composition of lactating Holstein dairy cows

Sixteen multiparous Holstein cows were used to determine the effects of 2-hydroxy-4-(methylthio) butanoic acid isopropyl ester (HMBi: 0 vs. 1.26 g/kg of total ration dry matter (DM) and dietary crude protein (CP) concentration [14.7% (low) vs. 16.9% (standard), DM basis] on milk yield and composition using a replicated 4 x 4 Latin square design experiment with 4-wk periods. Cows were fed ad libitum a total mixed ration with a 1: 1 forage-to-concentrate ratio (DM basis), and diets provided an estimated 6.71 and 1.86% lysine and methionine, respectively, in metabolizable protein for the low-protein diet and 6.74 and 1.82% in the standard protein diet. Dry matter intake, milk yield, and composition were measured during wk 4 of each period. There were no effects on DM intake, which averaged 24.7 kg/d. There was an interaction between dietary CP and HMBi for milk yield and 3.5% fat-corrected milk (FCM). Feeding HMBi decreased milk and FCM yield when fed with the low-CP diet but did not affect milk or FCM yield when fed with the standard CP diet. Feeding HMBi increased milk protein concentration regardless of diet CP concentration and increased milk protein yield when added to the standard CP diet but not the low-CP diet. The positive effect of HMBi on milk protein yield was only observed at the standard level of dietary CP, suggesting other factors limited the response to HMBi when dietary protein supply was restricted.

The effect of paddock size on the response of seasonal anoestrous ewes to the ram effect

Many aspects of the conditions required to maximize the ewe's response to ram introduction in the late anoestrous season remain unclear. The aim of this research was to determine whether grazing space allowances could influence the efficacy of the ram effect. In August 1995, at Reading (latitude 51degrees27'N), following a 3-month isolation period from rams, two groups of nulliparous Mule ewes, aged 15 months, were introduced to four rains in a low (12 ewes/ha; treatment L, n = 124) or in a high stocking rate (84 ewes/ha; treatment H, n = 126). From the beginning of August until the end of August oestrous behaviour was recorded by daily checks of mating marks on ewes. Rams were removed and in October all ewes were scanned (day 50) for pregnancy. No significant differences were found in the parameters investigated. Eighty-two percent of the L and 75.4% of the H ewes exhibited oestrus, with a pronounced peak on day 23 following ram introduction and a compact concentration in the 21-25-day period. The oestrous synchronisation rate in this 5-day period was 69.4 and 68.3%, respectively for L and H. The mean interval from ram introduction to oestrus was 23.17+/-2.4 days in L and 23.0+/-2.2 days in the H group. Conception rates were 84.3 and 87.4% for L and H groups, respectively. These results suggest that the response of anoestrous ewes to the introduction of rams was not affected by grazing space allowances and that yearling Mule ewes respond well to the ram effect in the late anoestrus season. (C) 2003 Elsevier Science B.V. All rights reserved.

The effect of wilting and soaking Eupatorium adenophorum on its digestibility in vitro and voluntary intake by goats

An experiment was conducted to determine what effect simple treatments might have on the voluntary intake by goats in Nepal of Eupatorium adenophorum, an invasive weed that is usually only consumed by goats to a very limited extent. Samples of E. adenophorum were collected and either untreated, soaked for 2 h or wilted for 2 h before being oven dried (60 degrees C) and ground. Soaking and wilting had little effect on the chemical composition of E. adenophorum, but did increase (P=0.036) its in vitro organic matter degradability, by approximately 8%. The short-term intake rate (STIR) of treated and untreated E. adenophorum was then estimated with eight goats. Soaking time (from 2 to 24 h) was not related to STIR (r = -0.111, P=0.198), but the time E. adenophorum was left to wilt (from 2 to 48h), was positively related to STIR (r=0.521, P<0.001), with values of STIR (g dry matter/min kg goat liveweight(0.75)) being 0.405, 0.649,1.058, S.E.M. 0.088 for E. adenophorum, that had been wilted for 0, 24 and 48 h respectively (P<0.001). Liveweight change of goats and voluntary intake of E. adenophorum by goats was then estimated with 24 goats. E. adenophorum was fed either unwilted, or wilted for 24 or 48 h. It was fed as the sole forage or as a 3:1 mixture (dry matter basis) with Ficus cunia. There was a linear (P<0.001) and quadratic (P<0.01) increase in the intake of total forage and E. adenophorum with wilting time of E. adenophorum. Offering Ficus cunia increased total forage intake, but decreased E. adenophorum intake (P<0.05). After four weeks, there was virtually no change in goat liveweight and no significant difference between treatments. The results suggest that wilting E adenophorum for 24 h could increase its intake by goats, and thereby increase its usefulness, as a potential source of forage in the dry season of Nepal, when forage scarcity is a common constraint to livestock production. (C) 2007 Elsevier B.V. All rights reserved.

Effect of prolonged intravenous glucose and essential amino acid infusion on nitrogen balance, muscle protein degradation and ubiquitin-conjugating enzyme gene expression in calves

Background: Intravenous infusions of glucose and amino acids increase both nitrogen balance and muscle accretion. We hypothesised that co-infusion of glucose ( to stimulate insulin) and essential amino acids (EAA) would act additively to improve nitrogen balance by decreasing muscle protein degradation in association with alterations in muscle expression of components of the ubiquitin-proteasome proteolytic pathway. Methods: We examined the effect of a 5 day intravenous infusions of saline, glucose, EAA and glucose + EAA, on urinary nitrogen excretion and muscle protein degradation. We carried out the study in 6 restrained calves since ruminants offer the advantage that muscle protein degradation can be assessed by excretion of 3 methyl-histidine and multiple muscle biopsies can be taken from the same animal. On the final day of infusion blood samples were taken for hormone and metabolite measurement and muscle biopsies for expression of ubiquitin, the 14-kDa E2 ubiquitin conjugating enzyme, and proteasome sub-units C2 and C8. Results: On day 5 of glucose infusion, plasma glucose, insulin and IGF-1 concentrations were increased while urea nitrogen excretion and myofibrillar protein degradation was decreased. Co-infusion of glucose + EAA prevented the loss of urinary nitrogen observed with EAA infusions alone and enhanced the increase in plasma IGF-1 concentration but there was no synergistic effect of glucose + EAA on the decrease in myofibrillar protein degradation. Muscle mRNA expression of the ubiquitin conjugating enzyme, 14-kDa E2 and proteasome sub-unit C2 were significantly decreased, after glucose but not amino acid infusions, and there was no further response to the combined infusions of glucose + EAA. Conclusion: Prolonged glucose infusion decreases myofibrillar protein degradation, prevents the excretion of infused EAA, and acts additively with EAA to increase plasma IGF-1 and improve net nitrogen balance. There was no evidence of synergistic effects between glucose + EAA infusion on muscle protein degradation or expression of components of the ubiquitin-proteasome proteolytic pathway.