Nerve cells in the hypothalamus make chemicals that control the release of hormones secreted from the pituitary gland. The hypothalamus gathers information sensed by the brain such as the surrounding temperature, light exposure, and feelings and sends it to the pituitary.
This information influences the hormones that the pituitary makes and releases. Pituitary: The pituitary puh-TOO-uh-ter-ee gland is at the base of the brain, and is no bigger than a pea. Despite its small size, the pituitary is often called the "master gland. The pituitary also secretes endorphins en-DOR-fins , chemicals that act on the nervous system and reduce feelings of pain. The pituitary also secretes hormones that signal the reproductive organs to make sex hormones.
The pituitary gland also controls ovulation and the menstrual cycle in women. Thyroid: The thyroid THY-royd is in the front part of the lower neck. It's shaped like a bow tie or butterfly. These hormones control the rate at which cells burn fuels from food to make energy. The more thyroid hormone there is in the bloodstream, the faster chemical reactions happen in the body. Thyroid hormones are important because they help kids' and teens' bones grow and develop, and they also play a role in the development of the brain and nervous system.
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As a consequence, some processes such as immune response mechanisms may be affected or delayed Kaattari and Tripp, ; Padgett and Glaser, ; Kudielka and Kirschbaum, For instance, hypothalamic CRH may act as an anti-inflammatory via stimulation of glucocorticoids and catecholamines; peripheral CRH acts as pro-inflammatory through direct action on immune cells Karalis et al. Besides, ACTH has been reported to present immunoreactive activity in the thymus of goldfish Carassius auratus Ottaviani et al. In addition, the expression of some immune genes in the central nervous system has been reported, and this suggests a potential cross-interaction between brain immune and neuroendocrine systems Metz et al.
Vaccination is the most effective method used nowadays in aquaculture to prevent diseases caused by pathogens Plant and LaPatra, Available data indicates that 2 h after Vibrio anguillarum bacterin exposure, the expression of both pro- and anti-inflammatory genes increase in gilthead seabream Sparus aurata head kidney primary cell culture Khansari et al. Therefore, these previous results demonstrate that a non-specific immune response is elicited in immune tissues of fish shortly after vaccination.
Also, serum or tissue antibodies such as immunoglobulin M and immunoglobulin T will increase at long-term after vaccination Lamers et al. Similarly, the phagocytic activity of head kidney leucocytes isolated from turbot Scophthalmus maximus L. There are several vaccine delivery methods, including oral, immersion and injection, of which injection often shows better protection Plant and LaPatra, However, the injection procedure can produce adverse reactions due to stress Hastein et al.
Nevertheless, few data is available regarding the effects of vaccine on the Hypothalamus-Pituitary-Interrenal HPI axis at the brain and pituitary level. In a previous study of our research group, it has been shown that bacterin could elicit immune responses in cultured pituitary cells of rainbow trout Oncorhynchus mykiss Liu et al. Taking all the above into consideration, the goal of the present study was to investigate the effect of vaccine in both brain and pituitary through different vaccination routes.
We hypothesized that: 1 bath vaccination might evoke a significant stress response of the central neuroendocrine organs of fish; and 2 the bacterin vaccines can induce both a stress and immune response in brain and pituitary. To test these hypotheses, two experiments were performed: In the first experiment, S. In the second experiment, we tested whether a vaccine administered through intraperitoneal injection was able to elicit a central stress response. It was also evaluated whether brain and pituitary showed a significant immune response.
As fish were taken out of the water for the injection, the responses to vaccination were tested against the air-exposed mock group, thus allowing consistent comparisons with the air exposure group from the first experiment. The air exposure stressor was selected for two reasons. One, because this is a previously used and validated type of stressor related to hypoxia or anoxia experiments Skrzynska et al. Second, because we wanted to differentiate the response of the vaccine itself compared to the response induced by a non-biotic physical stressor. Two batches of gilthead seabream During this period, no clinical signals of disease, malformation or injuries were observed, nor altered behavior.
The composition consists of inactivated V. The second stressor, air exposure, consisted in 3 min out of the water. There were two replicate tanks in each group. It is worth to mention that vaccination was performed 24 h before air-exposure stress since the preliminary result in systemic immune organs did not show any significant alteration by vaccine at early time of vaccination data not shown. Fish were sampled after 1, 6 and 24 h.
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Fish were anesthetized by an overdose of tricaine methanosulphonate MS and the blood from each fish was quickly collected from the caudal vein by using a heparinized 2 mL syringe. Before the start of the experiment, a total of 36 fish were randomly divided into 2 groups with two replicate tanks per group as for mock injection and vaccination, and these fish were acclimatized for another 5 days in L water circular tanks. During this period, water parameters and rearing conditions were kept the same as mentioned above.
After the injection, fish were immediately returned to the corresponding experimental tanks. The whole operation lasted less than 3 min. Fish from both mock injection and vaccination groups were sampled at 1, 6, and 24 h post injection, and blood, pituitary and brain of 6 fish from each group at each sampling time point were collected. In brief, fish was anesthetized by an overdose of MS, the blood of each fish was quickly collected from the caudal vein by using a 5 mL syringe, which was pre-rinsed with lithium heparin Deltalab, Spain , and then transferred to a clean tube with one drop of lithium heparin.
After fast blood collection, the pituitary gland and brain of each fish were excised, immediately frozen in liquid nitrogen and stored under o C until use. The composition consists of inactivated L. These Elisa kits used in the present study showed sensitivities of about 1. Seabream plasma cortisol levels were measured by radioimmunoassay RIA as described by Rotllant et al. A melting curve analysis was carried out after the completion of RT-qPCR to verify no non-specific amplification. The reference genes 18S and RPL27 were used for normalization. The quantification was performed according to Pfaffl method Pfaffl, and corrected for the efficiency of each primer set.
Value for each experimental condition was expressed as normalized relative expression, calculated in relation to the values of control group and normalized against those of the reference gene 18S. The amplification efficiency and product size are listed in Table 1. Six biological replicates with two technical replicates were performed for the qPCR analysis. For the first experiment the statistical package for social science SPSS, v20 software was used for the analysis.
The Generalized Linear Model GzLM was utilized considering the stressors and time dynamics as a two between-subjects factor. This model is a more flexible statistical tool than the standard general linear model GLM in terms of types of distribution and different covariance structure of the repeated measures does not require homogeneity of variance and it admits missing values. After the main analysis, appropriate pairwise comparisons were carried out. Figure 1 shows the levels of plasma cortisol after both stressors and its combination at the respective time points.
Cortisol did not show any significant response after bath vaccine treatment compared to air exposure stress, which showed a classical acute response dynamics with a peak at 1 h, still significantly higher at 6 h, followed by further recovery of basal values at 24 h. The dynamics of the vaccine plus air exposure was similar to that observed after air exposure, although the recovery took place later on, indicating that the air exposure stressor was predominant in the cortisol response.
Figure 1. Plasma cortisol concentration in gilthead seabream Sparus aurata L. Significant differences are indicated by numbers in air exposure group and by capital letters in vaccine plus air exposure group.
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Regarding the response of the analyzed stress-related genes in the brain, no relevant changes were observed after bath vaccine treatment, except for a decrease of crhbp at 1 h, whereas air exposure showed significant increases in crh, crhbp , and gr and a significant down-regulation of trh.
When both stressors were applied, trh was maintained down-regulated at 6 h and only crhbp increased significantly Figure 2. Figure 2. Significant differences are indicated by capital letters in vaccine group, by numbers in air exposure group, and by lowercase letters vaccine plus air exposure. In the pituitary, bath vaccination showed a differential induction of pomc genes at short time 1 h whereas at 6 h prl and gh showed significant increases. Air exposure increased the expression of pomcb, gr and sl1 at 1 h, and gr at 6 h.
After applying both stressors only slight changes were detected as for the reductions of pomcb at 6 h and sl1 at 1 h, and the increase of sl1 at 6 h Figure 3. Figure 3. Plasma cortisol values significantly raised by 2. The differential cortisol increase of the vaccine-injected fish compared to the mock-injected fish was apparent at all time points. Regarding time course, both injected vaccine and mock groups presented the same cortisol dynamics, i. However, the vaccinated group showed higher levels at either time compared to mock-injected group.
The vaccine injected group also showed higher resistance to recovery at 24 h. Regarding CRH or ACTH, no alteration of plasma content in either group mock or vaccination was observed in none of different time points assessed 1, 6, and 24 h post injection Figure 4.
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Figure 4. In the second experiment, the expression of crh was almost unchanged at 1, 6, and 24 h post vaccination. Transcript of gr was not altered at 1 or 24 h, while there was a slight but significant up-regulation at 6 h post vaccination in the mock group. A similar trend can be observed in the heat shock proteins HSP hsp70 and hsp90 in which a significant increase was also observed at the same time point 6 h. As a whole, few changes were observed in brain genes, and the changes were higher in mock-injected fish than in vaccine-injected fish Figure 5.
Figure 5. Similar than with the bath vaccine, the expression of stress genes in the pituitary showed a different pattern in which one gene, CRH binding protein crhbp , substantially increased its expression up to 15 fold at 6 h or up to 7 fold after 24 h after vaccine injection. These increases contrast with the mock injected fish in which the increase was moderate between 3 and 5 fold , though showing the same dynamics. A similar significant trend, but more moderate over two fold increases , was observed for hsp70 but not for hsp The rest of the genes assessed, although showing some variations, did not change significantly their expression except for gr in which a significant down-regulation was observed at 6 h Figure 6.
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In the second experiment it was intended to determine whether an injected vaccine induced immune gene expression changes in brain and pituitary other than in stress-related genes. The results showed a very clear picture as not only some cytokines increased their expression but the level of induction was very strong. Similarly, cox2 was significantly up-regulated by 5. Other pro-inflammatory gene transcripts such as il6 raised significantly only 1. Nevertheless, the mean values showed a non-significant but apparent increasing trend Figure 7.
The expression of lysozyme gene lys significantly increased in the brain of seabream at 6 h post vaccination, although with just 1. The transcript for the complement C3 component gene c3 showed a significant increase at 1 h post vaccination, by 2. Figure 7.
Regarding pituitary, the expression of genes related to the immune responses are shown in Figure 8. A similar alteration trend was observed for the expression of cox2 after vaccination, however, with less intensity. The cox2 transcript raised by 4. Compared to the mock injection, lys showed comparable levels at 1 h post injection, while it was distinctly up-regulated by Figure 8.
The combined results from the two vaccination experiments indicate that fish perceives the vaccine as a stressor but at a limited extent. Thus, bath vaccine did not induce plasma cortisol rise, while injection vaccine did produce a differential cortisol response compared to mock injection. Therefore, in terms of plasma hormones, it seems that fish would not perceive vaccines as primary stressors stimulating the hierarchical activation of HPI axis, although it would indirectly activate cortisol release in the case of vaccine injection linked to the air exposition period during the injection procedure.
In both experiments cortisol presented an acute response dynamics, peaking at 1 or 6 h and recovering at 24 h, in agreement with the studies previously reported for this species after subjecting seabream to acute stressors such as air exposure Arends et al. Similar increases of cortisol concentration in rainbow trout treated by vaccine injection have been previously reported as well Funk et al.
However, in other works in which higher doses of bacteria were administered to Eleginops maclovinus or S.
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Our results also suggest that vaccines do not clearly activate the response of brain stress genes during the first hours. Thus, neither crh nor crhbp or gr showed relevant modulation after bath vaccine and only slight changes were observed in hsp and gr after injection. Therefore, this suggests that vaccine did not activate the central stress gene response unless a physical stressor air exposure was included, as observed in the vaccine plus air exposure groups. This agrees with the previously reported response of S.
On the contrary, in the pituitary, vaccine did induce the gene expression of stress-related hormones like prl, gh at 6 h and pomca and pomcb peptides at 24 h. Therefore, it seems that the pituitary was more sensitive than brain to immune stimulation, although at later time points 6 and 24 h.
This may indicate, other than a higher sensitivity, that the pituitary stimulation could be not a direct effect, but resulting from the interaction through biological messengers such as cytokines. This would be supported by the fact that both brain and pituitary showed a robust pro-inflammatory cytokine response to vaccine see Figures 7 , 8. Getting Started. Surgery Guide. Send Us a Message.
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