Comparison of rolled, NaOH treated and ensiled wheat grain in dairy cattle diets

The effect of three wheat grain treatments on the zootechnical performance (feed intake, milk production and milk fat and protein production) of dairy cattle was studied with 18 Holstein cows in each of two feeding trials (both a 3 × 3 Latin square design). Rolled wheat, NaOH treated wheat and immature wheat ensiled with Graintona-Plus (Vee-lik, Zuienkerke, Belgium) were compared in a ration with fixed DM proportions of wheat, maize silage and prewilted grass silage. The nutritive value of the three wheat treatments was derived from in situ rumen degradability studies and from in vivo digestibility experiments.

In situ rumen incubations showed that NaOH treatment strongly decreased protein and starch degradability of the wheat. Due to that importantly decreased starch degradability NaOH treated wheat may be a source of starch with a reduced risk of rumen acidosis as compared to rolled wheat. On the other hand, due to the ensiling, immature wheat had a very high washable starch fraction, despite the Graintona-Plus treatment. The protein value differed importantly, with the DPI (true protein digested in the small intestine; [Tamminga, S., van Straalen, W.M., Subnel, A.P.J., Meijer, R.G.M., Steg, A., Wever, C.J.G., Blok, M.C., 1994. The Dutch protein evaluation system: the DVE/OEB-system. Livest. Prod. Sci. 40, 139–155]) value being 103, 125 and 76 g/kg DM for rolled, NaOH treated and ensiled wheat, respectively. The RDPB (degraded protein balance; [Tamminga, S., van Straalen, W.M., Subnel, A.P.J., Meijer, R.G.M., Steg, A., Wever, C.J.G., Blok, M.C., 1994. The Dutch protein evaluation system: the DVE/OEB-system. Livest. Prod. Sci. 40, 139–155]) value of the NaOH treated wheat was − 51 vs. − 37 and− 33 g/kg DM for rolled and ensiled wheat. In vivo digestibility trials with Holstein cows indicated that the net energy value of the wheat treatments was not significantly different.

In Trial 1, all diets were formulated to supply 105% of the requirements of DPI and net energy lactation (NEL; [Van Es, A.J.H., 1978. Feed evaluation for ruminants: I. The system in use from May 1977 onwards in the Netherlands. Livest. Prod. Sci. 5, 331–345]) and a safe RDPB level (± 200 g/day). On average 4.6 kg wheat DM was fed daily. Following trial 1, trial 2 was carried out, with the maize silage, grass silage and wheat fed in the same proportions as in trial 1, but fed ad libitum, to compare the intake level of the three diets. In both trials, the NaOH treated wheat resulted in the best performance with importantly increased milk yield and the highest yield of fat and protein corrected milk. Ensiled immature wheat resulted in a significant lower milk production and milk protein yield and in a higher milk fat content. In trial 1, DM-intake of the three groups was comparable due to the experimental design. In trial 2 intake of the cows fed NaOH treated wheat was significantly higher (23.5 vs. 21.1 kg/day for the other two groups).     

不同品种青贮玉米生产性能及发酵后营养价值评价

试验旨在筛选适合宁夏南部冷凉地区推广种植的青贮玉米品种,利用灰色关联理论对17个青贮玉米品种的株高、鲜草产量等主要农艺性状、生产性能及发酵营养指标进行综合评价.结果 显示,不同品种的各单项指标表现不同,且各单一最优指标并不聚集于同一品种中.研究表明,桂青贮1号、宁单34、昊玉501、金岭10、京科青贮516的灰色关联度...     

A major outbreak of botulism in cattle being fed ensiled poultry litter

Eighty of a group of 150 housed beef cattle showed classical signs of botulism after eating a batch of ensiled poultry litter. Sixty-eight of the animals died and Clostridium botulinum type C toxin was detected in 18 of 22 sera examined. C botulinum organisms were isolated from the ensiled litter and type C toxin was demonstrated in samples of decomposed poultry carcases present in the litter. This outbreak of bovine botulism was the most serious to have been recorded in Europe and was the first associated with feeding ensiled poultry litter.     

Feeding value of urea molasses-treated wheat straw ensiled with fresh cattle manure for growing crossbred cattle calves

The study was carried out to evaluate the influence of urea plus molasses-treated wheat straw (WS) ensiled with cattle manure (CM) on nutrients intake, their digestibilities, and growth performance of crossbred (Sahiwal × Holstein Friesian) cattle calves. The CM was mixed with ground WS in a ratio of 30:70 on dry matter (DM) basis. The WS–CM mixture treated with urea (4% DM) and molasses (4% DM) was allowed to ferment for 40 days in a cemented pit. Four iso-nitrogenous and iso-energetic fermented wheat straw (FWS)-based experimental diets were formulated. The FWS0, FWS20, FWS30, and FWS40 diets contained 0%, 20%, 30%, and 40% FWS, respectively. Twenty calves (9–10 months of age) were randomly allocated to four dietary treatments in a randomized complete block design, five in each group. Increasing trends for DM, organic matter, crude protein, and neutral detergent fiber intakes by calves were observed with increasing dietary FWS level. Weight gain was significantly different among calves fed different levels of FWS. The highest weight gain (491.8 g/day) was observed in calves fed FWS40 diet, while calves fed FWS0 and FWS20 diets gained 350.0 and 449.6 g/day, respectively. The results from this study imply that the FWS can be added up to 30% in the diet of growing crossbred calves without any detrimental effect on their performance.     

Immune responses of infected and vaccinated Hereford cattle to antigens of the cattle tick, Boophilus microplus

Responses of infested and vaccinated Hereford cattle to Boophilus microplus antigens were measured by enzyme-linked immunosorbent assay (ELISA), lymphocyte blastogenesis assay (LBA) and intradermal skin tests. Responses against soluble salivary gland extracts (SGS), salivary gland membrane (SGM), soluble gut extracts (GS), gut membrane (GM), soluble larval extracts (LS) and larval membrane (LM) antigens were tested. In one experiment, cattle infested with up to 160,000 ticks had positive cellular responses to SGS and significant antibodies against LM, GM, SGM, and SGS. Cellular responses to Concanavalin A were not depressed following infestation. Cattle vaccinated with GM, using Quil A as adjuvant, had positive cellular responses to gut and salivary gland antigens and significant antibody responses to all antigens tested. The antibody levels of vaccinated cattle were significantly higher than the antibody levels of infested cattle (P less than 0.05). In a second experiment, immune responses of cattle infested with 40,000 ticks were studied during 38 days. Cellular responses in LBA to several tick antigens were transiently elevated and significant levels of antibody were measured against LM, GM, SGM and SGS, from day 25 (P less than 0.05). Infested cattle had positive skin reactions following intradermal injection of larval and adult tick antigens (P less than 0.05).     

Behavioral, adrenal, immune, and productive responses of lactating ewes to regrouping and relocation.

The experiment involved 40 mid-lactating Comisana ewes. The animals were housed in straw-bedded pens and assigned to a control group (n = 10) and two test groups of 15 subjects each. Control ewes were never moved from their pen and peers throughout the experiment period, whereas test animals were subjected to either regrouping (RG) or regrouping and relocation (RGRL) three times at weekly intervals (d 7, 14, and 21 of the trial). Three behavioral recordings were conducted after each social and pen exchange over a 6-h period (1000 to 1600). Animals from groups RG and RGRL were bled immediately before each regrouping and relocation and 15 and 60 min afterward. The phytohemagglutinin skin test was performed at d 9, 16, and 23. Milk yield was recorded when the ewes were mixed and moved, and before and after each mixing and moving procedure. Individual milk samples were analyzed for composition, renneting parameters, and bacteriological characteristics; samples with more than 10(6) somatic cells/mL were cultured for mastitis-related pathogens. Control ewes spent more time lying than groups RG (P < 0.001) and RGRL (P < 0.01) and less time in ambulatory activities than RGRL sheep (P < 0.05). Moved and remaining ewes showed a higher number of aggressive interactions than control subjects (P < 0.001 and P < 0.01, respectively). Control sheep showed the highest cell-mediated immune response compared with both moved and mixed ewes (P < 0.001 and P < 0.01, respectively). The RGRL ewes gave lower yields of milk than did the control ewes on the days after the first (P < 0.05) and the second (P < 0.01) change of pen and peers. The RGRL ewes also had a lower fat content (P < 0.05) than control ewes in the milk yielded on the day of the first regrouping and relocation and a lower milk protein content (P < 0.05) on the following day. The RG treatment resulted in the ewes yielding milk with a lower fat content (P < 0.01) on the day of each mixing procedure as compared with controls and in a decreased milk protein content on the days after the first (P < 0.05) and the second regrouping (P < 0.01). One case of subclinical mastitis was recorded in both the RG and RGRL groups, whereas no cases were detected in control group. These findings suggest that regrouping and relocation may cause increased aggression, altered immune responses, and short-term effects on the production performance of lactating ewes. These practices should be performed by attempting to minimize their impact on animal welfare.     

Behavioral and physiological responses to stabling in naive horses

The purpose of this study was to investigate the response of horses to confinement and isolation in a stable (indoor individual housing) for the first time using behavioral indices, heart rate, and salivary cortisol concentration. Six naive 2-year-old Australian Stock Horse fillies were examined at 4-hour intervals over 24 hours in an outdoor group paddock followed by 24 hours in indoor individual housing. Behavioral observations and scores and heart rates were recorded and saliva samples were taken at each interval. During stabling, all horses became agitated and demonstrated increased vocalization and movement. Behavioral scores were significantly higher in the indoor individual housing (P < .001). No significant difference in heart rates between the two environments was detected. Mean salivary cortisol did not increase significantly (2 ng/mL ± 1.4 ng/mL in outdoor group paddock vs 2.5 mL ± 1.2 ng/mL in indoor individual housing). No diurnal rhythm in salivary cortisol was evident in either the outdoor group paddock or indoor individual housing. The results of this study highlight that a combination of behavioral and physiological measures allow better understanding of stress, where one measurement may be misleading. First time stabling of horses elicited marked behavioral responses indicative of stress that were not reflected in increased heart rates or salivary cortisol concentrations. The lack of a diurnal cortisol rhythm and the comparatively high basal cortisol concentrations found in the outdoor group paddock environment may imply that the fillies were already stressed; therefore, stabling did not cause further aberrations detectable by salivary cortisol analysis.     

Methane emission by Nellore cattle subjected to different nutritional plans

The objective was to evaluate the emission of enteric methane by Nellore cattle subjected to different nutritional plans, as well as the intake and digestibility of nutrients from the diets supplied. Forty-seven animals in a confinement system (feedlot) were fed a corn silage-based diet for 35 days. Afterwards, these animals were evaluated in a grazing system during the rainy periods, in Urochloa brizantha cv Marandu paddocks, for 44 days. Chromic oxide was used as external marker to estimate the fecal production of animals. Samples of feces, ingredients, and pasture were collected and sampled for subsequent chemical analyses. The SF₆ tracer gas technique was adopted to quantify the methane gas emitted by the animals. The experimental design was completely randomized, using procedure GLM of software SAS (9.2), including the fixed effects of sex and nutritional plan and the linear effect of the co-variable weights. Means were analyzed by Tukey’s test at 5 % probability. The intakes of total dry matter and organic matter were greater for the animals subjected to the feedlot diet (P?P?P?4/day (104.01 g) by the animals fed the feedlot diet (P?4/CEB) and methane emission per dry matter consumed were lower in relation to the grazing treatment (3.75 vs 4.23 % and 11.67 vs 15.71 g/kg). The better quality of the diet in the feedlot promoted lower energy losses as methane.     

Dynamic responses of cattle to thermal heat loads

The focal point of this limited review is bioenergetic research conducted in the Biological Engineering Research Unit at the U.S. Meat Animal Research Center (MARC), using recently developed instrumentation and analytical techniques. The dynamics of observed thermoregulatory responses in cattle to thermal heat load challenges are explored, with an emphasis on physiological and behavioral parameters of body temperature, respiration rate, and feed intake. Observations of body temperature, especially tympanic temperature, have shown hot environments to cause phase shifts, increased amplitude, and increased means for diurnal rhythms. Fractal analysis of body temperature records obtained at 2- to 10-min intervals has been found to be robust for objectively differentiating among responses of cattle in cool to hot environments, and it indicates a stress threshold of approximately 25 degrees C (coincident with declining feed intake). Other analyses determined a 21 degrees C threshold for increased respiration rate. The reported observations and analyses provide further understanding of how and why the animals respond to environmental challenges, an understanding that is necessary for refining performance models and developing energetic and thermoregulatory models. The dynamic responses are discussed in the context of establishing criteria for proactive environmental management for cattle during hot weather, using heat waves as an example.     

Regulation of IgA responses in cattle, humans and mice

Secretory IgA (SIgA) constitutes the largest component of the humoral immune system of the body with gram quantities of this isotype produced by mammals on a daily basis. Secretory IgA (SIgA) antibodies function by both blocking pathogen/commensal entry at mucosal surfaces and virus neutralization. Several pathways of induction of IgA responses have been described which depend on T cells (T cell dependent or TD) pathways or are independent of T cells (T-independent or TI) and are mediated by dendritic cells (DCs) and/or epithelial cells. Many elements of IgA regulation readily cross species barriers (adjuvants, soluble and cognate factors) and are highly conserved whereas other pathways may be more specific to any given species and must be evaluated. Regulation of IgA production in cattle is not completely understood and thus we have focused in part on highly conserved factors such as transforming growth factor beta, Type I and Type 2 interferons, neuropeptides which interdigitate mucosal tissues (vasoactive intestinal peptide or VIP), and a small peptide (IgA inducing peptide or IGIP) which can serve as targets for modulation and increasing SIgA virus-specific antibodies. We have evaluated the potential utility of modulating these factors in vitro in regulation of qualitative aspects of antibodies of the IgM, IgG and IgA isotypes at mucosal surfaces and in secretions of the upper and lower respiratory tract to a virus of economic and public health importance, foot and mouth disease virus (FMDV). IgA responses in cattle are essential for host defense in response to various infectious agents. In cattle, IgA is not released into the colostrum, as is the case for other mammals but only IgG1 is selectively transported. In previous studies in cattle, IgA has been shown to be regulated by several cytokines including IFN-gamma, Type I interferons such as IFN-alpha and IFN-tau, transforming growth factor beta, IgA inducing peptide and other potential factors such as APRIL and BlyS which have not yet been fully evaluated in cattle. Many of these factors, namely TGF-beta and Type I interferons block cell cycle progression which is an essential component of Ig class switching and thus these factors require additional regulatory factors such as IL-2 to drive cells through cell cycle resulting in class switch recombination. Among these factors, IgA inducing peptide was originally identified from a bovine gut associated lymphoid tissue expression library and is highly conserved in pigs and humans at >90% at the amino acid level. The factor is regulated differently in various species but is consistently produced by dendritic cells.     

Behavioral responses of turkeys subjected to different climatic conditions

I was evaluated the effect of seven different combinations of temperature, air velocity, and relative air humidity on the frequency and duration of eating, drinking, resting, cannibalism, dust bathing, scratching, ground pecking, shivering, and stretching behaviors of turkeys at three different ages. The combinations tested of temperature, relative air humidity, and air velocity were, respectively: 1 (22 °C, 50%, 1 m/s); 2 (26.2 °C, 73.2%, 0.45 m/s); 3 (26.6 °C, 71.2%, 1 m/s); 4 (28.9 °C, 72%, 1.4 m/s); 5 (31.1 °C, 85%, 0.45 m/s); 6 (34.1 °C, 82.1%, 1 m/s); and 7 (34.4 °C, 82.1%, 1.4 m/s) for three ages of birds (61, 96, and 131 days of age). Seven birds were housed per pen, at a density of 3 males/m², totaling 147 birds in the entire experiment. Each combination was applied for 5 days. The data were analyzed considering the number of times the bird performed the behavior and the time it performed (in seconds). Each pen was considered a repetition. A comparison of the medians was used to compare the treatments by each age. The results showed that young birds were more likely to suffer from the combination of low temperature and high air velocity, reducing their frequency of normal behaviors. Increased humidity at a low temperature raised the frequency of scratching, shivering, and cannibalism behaviors leading to poorer bird welfare. It is recommended that the temperature, relative air humidity, and air velocity combination of 26.6 °C; 71.2%; and 1 m/s, respectively, for young birds, and 22 °C; 50%; and 1 m/s, respectively, for older birds should be used.

     

Adrenocortical responses to tumor necrosis factor-alpha and interferon-gamma in cattle

The responses of plasma cortisol and adrenocorticotropic hormone (ACTH) were examined to intravenous injection of recombinant bovine tumor necrosis factor-alpha (TNF-alpha) and interferon-gamma (INF-gamma) in Holstein cows. INF-gamma induced dose-dependent rises in the plasma levels of both cortisol and ACTH, while TNF-alpha induced comparable plasma cortisol responses with much smaller rises in plasma ACTH. The results suggest a direct stimulatory action of TNF-alpha on cortisol secretion from the adrenal gland in cattle.     

Fermentation characteristics and nutritional value of ruminal contents and blood ensiled with untreated or sodium hydroxide-treated wheat straw.

Bovine ruminal contents and blood were ensiled with wheat straw, either untreated or treated with 4% NaOH (DM basis), with and without addition of 5% dry molasses. Proportions of ruminal contents, blood, and straw were 45:15:40, wet basis, respectively. Wheat straw also was ensiled alone, and water was added to attain 43% DM. After 60 d, all silages had a desirable aroma. The pH of the ensiled mixtures of ruminal contents, blood, and untreated straw was 4.43 and 4.56, with and without molasses, respectively. Values for silages containing treated straw, with and without molasses, were 5.95 and 7.37, respectively. All silages had substantial levels of lactic acid (2.4 to 4.9%, DM basis). Addition of molasses increased (P less than .01) lactic acid concentrations. In a metabolism trial, 36 wethers were fed a 50% orchardgrass hay basal diet alone or supplemented with ensiled materials (1:1 ratio, DM basis) such that final diets had about 7 and 4% of their DM from ruminal contents and blood, respectively. Among the diets containing the ensiled mixtures, digestibilities of DM, OM, energy, NDF, ADF, cellulose, and hemicellulose were 6 to 20 percentage units higher (P less than .01) for silages containing treated vs untreated straw. Addition of molasses increased apparent digestibility only of CP and only in sheep fed ensiled mixtures with treated straw (interaction of straw treatment x molasses addition, P less than .05). Nitrogen balance was increased (P less than .05) from 1.6 g/d to 2.9 g/d by NaOH treatment of straw. Metabolizable energy was higher (P less than .05) for diets containing silages with NaOH-treated straw.(ABSTRACT TRUNCATED AT 250 WORDS)     

Modelling responses to nutritional, endocrine and genetic strategies to increase fertility in the UK dairy herd

The United Kingdom, as in most countries using intensive dairy management programmes, is facing serious challenges in terms of dairy cow fertility, as highlighted by a rapidly increasing calving interval (CI). A mechanistic, mathematical model is described that predicts the size of the future national dairy herd required to supply domestic requirements and its inherent sustainability in terms of production of replacement female numbers. The results from the model suggest that continuing use of current management strategies may result in the national dairy herd being unsustainable due to increasing CI and reduced fertility in as few as 10 years. Adoption of nutritional, endocrine and genetic techniques that increase fertility can effectively and rapidly reverse this trend and reduce the required size of the national herd, thereby reducing methane emissions from dairy production.     

Physiological responses of Bos taurus and Bos indicus cattle to prolonged, continuous heat and humidity

Two experiments were conducted to investigate the physiological responses of Bos taurus (Angus cross, n = 6) and Bos indicus (Brahman, n = 6) cattle to prolonged heat and humidity, as can occur during live export by sea. Each experiment was carried out in climate-controlled rooms, where heifers were exposed to 15 d of sustained heat and humidity. The treatment was designed to be representative of a long-haul, live-export voyage leaving a southern Australian winter and traveling to a Middle Eastern summer. Wet bulb temperature (WBT) was used to give a combined measure of dry bulb temperature and relative humidity and was increased over several days, culminating in 5 d at 32 degrees C WBT between d 7 and 11. By d 11, the respiratory rate and core body temperature increased (P < 0.001) compared with values at lower ambient temperature on d 1 and 2 when climate-controlled rooms were not operating. Feed intake of Bos taurus was reduced (P < 0.001) by d 11, whereas that of Bos indicus did not change (P = 0.14). Despite no diurnal variation in climatic conditions, core body temperature of both Bos taurus and Bos indicus continued to show a circadian amplitude of approximately 1 degrees C throughout the hottest period. This amplitude increased during the recovery period after heat was removed (up to 1.8 degrees C for Bos indicus and 1.6 degrees C for Bos taurus). Water intake for both Bos taurus and Bos indicus increased when WBT increased (P < 0.01 on d 11). Significant acid-base and blood electrolyte imbalances occurred in both Bos taurus and Bos indicus, with changes in Bos taurus being more substantial and prolonged. The increase in respiratory rate coincided with a decrease in the partial pressures of carbon dioxide and bicarbonate in venous blood. However, during the hottest period, average daily venous blood pH remained unchanged. When the heat load was reduced after d 11, the blood pH decreased, indicating metabolic acidosis. Blood pH declined from 7.44 to 7.36 for Bos taurus (P < 0.001) and from 7.44 to 7.38 for Bos indicus (P < 0.001). Other parameters measured include heart rate; packed cell volume; plasma and urine Na, K, and Cl; urine pH; and specific gravity. Our results suggest that Bos taurus cattle experience significant physiological changes during exposure to prolonged and continuous high heat and humidity, with alterations persisting for some days after the heat-stress conditions subside. Bos indicus experience similar but less pronounced physiological changes.     

Behavioral responses to two intranasal vaccine applicators in horses and ponies

OBJECTIVE: To evaluate behavioral compliance of horses and ponies with simulated intranasal vaccination and assess development of generalized aversion to veterinary manipulations. DESIGN: Clinical trial. ANIMALS: 28 light horse mares, 3 pony geldings, 2 light horse stallions, and 3 pony stallions that had a history of compliance with veterinary procedures. PROCEDURE: Behavioral compliance with 2 intranasal vaccine applicators was assessed. Compliance with standard physical examination procedures was assessed before and after a single experience with either of the applicators or a control manipulation to evaluate development of generalized aversion to veterinary manipulation. RESULTS: In all 30 horses, simulated intranasal vaccination or the control manipulation could be performed without problematic avoidance behavior, and simulated intranasal vaccination did not have any significant effect on duration of or compliance with a standardized physical examination that included manipulation of the ears, nose, and mouth. Results were similar for the 2 intranasal vaccine applicators, and no difference in compliance was seen between horses in which warm versus cold applicators were used. For 3 of the 6 ponies, substantial avoidance behavior was observed in association with simulated intranasal vaccination, and compliance with physical examination procedures decreased after simulated intranasal vaccination. CONCLUSIONS AND CLINICAL RELEVANCE: Although some compliance problems were seen with ponies, neither problems with compliance with simulated intranasal vaccination nor adverse effects on subsequent physical examination were identified in any of the horses. Further study is needed to understand factors involved in practitioner reports of aversion developing in association with intranasal vaccination.     

Production responses to internal parasite control in dairy cattle

A significant increase in milk production, averaging 164 litres per cow per lactation (a 4.8% increase), was seen after cows infected with gastrointestinal nematodes, paramphistomes and Fasciola hepatica were treated with broad-spectrum anthelmintics. Three hundred and ninety pairs of cows from eight herds with year-round calving were studied. One cow in each matched pair was given 7.5 mg/kg fenbendazole, 7.5 mg/kg levamisole hydrochloride and 15 mg/kg oxyclozanide in March, May and August of one year; the other cow in the pair received no anthelmintic. The number of nematode and trematode eggs was significantly decreased in the faeces of treated cows.     

Immune responses of cattle to experimental anti-Fasciola hepatica vaccines.

Fasciola hepatica infection of cattle and sheep is an important cause of clinical disease and production losses, and is controlled at present by a combination of chemotherapy and management measures. However, the prospects for the control of F. hepatica infection by vaccination are good, and we have previously shown substantial protection of cattle against experimental challenge infection following immunisation with a combination of the purified fluke-derived enzymes cathepsin L1 (CATL 1), cathepsin L2 (CATL 2) and fluke-derived Hb fraction (FHB). This and other recent studies have also demonstrated fundamental differences between protective and non-protective immune responses to liver fluke infection. In this present study we have further analysed the response of animals to liver fluke challenge following experimental vaccination. Calves were vaccinated with either CATL 2 plus FHB, or CATL 1 plus CATL 2. Partial protection against challenge infection was achieved in both vaccinated groups, with the greatest level of protection (55 per cent reduction in fluke burdens) recorded in the group vaccinated with CATL 1 plus CATL 2. This latter group also showed the greater level of lymphocyte proliferation and the greater production of gamma-INF in response to stimulation with fluke antigen in vitro following challenge. These results are significant in our attempts to characterise the elements within the immune response to vaccination which are protective.