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|Table of Contents
|Vol. 66, No. 3, 2006
Issue release date: August 2006
|Section title: Mini Review
Autoimmunity: Basic Mechanisms and Implications in Endocrine Diseases
Part IIBallotti S.a · Chiarelli F.b · de Martino M.a
aDepartment of Paediatrics, Anna Meyer Children's Hospital, University of Florence, Florence, and bDepartment of Paediatrics, University of Chieti, Chieti, Italy Corresponding Author
Prof. Maurizio de Martino, MD
Department of Paediatrics, University of Florence
via Luca Giordano, 13
IT–50132 Florence (Italy)
Tel. +39 0555 662 494, Fax +39 0555 703 80, E-Mail firstname.lastname@example.com
Regulation of the immune response to self-antigens is a complex process that involves maintaining self-tolerance while preserving the capacity to exert an effective immune response. The primary mechanism that leads to self-tolerance is central tolerance. However, potential pathogenic autoreactive lymphocytes are normally present in the periphery of all individuals. This suggests the existence of mechanisms of peripheral tolerance that prevent the initiation of autoimmune diseases by limiting the activation of autoreactive lymphocytes. If these mechanisms of peripheral tolerance are impaired, the autoreactive lymphocytes may be activated and autoimmune diseases can develop. Several processes are involved in the maintenance of peripheral tolerance: the active suppression mediated by regulatory T cell populations, the different maturation state of antigen-presenting cells presenting the autoantigen to autoreactive lymphocytes, inducing tolerance instead of cell activation, the characteristics of B cell populations. A deeper comprehension of these mechanisms may lead to important therapeutic applications, such as the development of cellular vaccines for organ-specific autoimmune diseases. In addition, autoimmunity does not always have pathological consequences, but may exert a protective function, as suggested by several observations on the beneficial role of autoreactive T cells in central nervous system injury.
© 2006 S. Karger AG, Basel
Regulation of immune responses to self-antigens is a complex process that involves maintaining self-tolerance while preserving the capacity to exert an effective immune response. Mechanisms of central tolerance are insufficient to control autoimmune events. Autoreactive lymphocytes are present in most normal individuals, but only 5% of the general population is affected by autoimmune diseases. This suggests the existence of multiple mechanisms of peripheral tolerance, whose function is to keep autoimmune phenomena under control.
In this second part the processes underlying peripheral tolerance have been analyzed focusing on the cellular mechanisms.
Regulatory T cells (Tregs) may be defined as T cells that inhibit effector T cell proliferation in vitro and in vivo[1]. Several cells with regulatory properties exist which may be included in two categories. Some Tregs arise after antigen exposure: Th2 cells; Th1 cells; interleukin (IL)-10 producing Tr1 cells; transforming growth factor-β (TGF-β) producing Th3 cells; CD8+ regulatory cells, and γδ T cells. Other Tregs spontaneously develop: CD4+CD25+ T cells; natural killer T cells, and probably γδ T cells (although they increase after antigen sensitization). A large overlap may exist among these cell subsets which exert their function as one or another subset accordingly to their activation stage or microenvironment [2,3,4].
The constitutive expression of IL-2 receptor α-chain (CD25) is likely to be the most specific Treg marker. CD4+CD25+ T cells represent up to 10% of the human CD4+ T lymphocytes. However, CD25 is not an exclusive Treg marker as its expression is induced on CD4+CD25– T cells during immune responses. Nevertheless, Tregs express CD25 at a higher level and more stably, while CD4+CD25– T cells lose CD25 when the immune response switches off [5,6,7]. Tregs constitutively also express adhesion molecules (LFA-1, CD44, ICAM-1, CD103, CCR4 and CCR8) which guarantee rapid recruitment to the inflammation site. Tregs additionally express CD152, co-stimulatory molecule cytotoxic T-lymphocyte antigen 4 (CTLA4) and the tumor necrosis factor (TNF) super family member glucocorticoid-induced TNF receptor (GITR) family-related protein TNFRSF18. These molecules classically upregulate after T cell receptor (TCR) activation. The partially activated phenotype of Tregs may depend on the ongoing engagement of their TCRs by autoantigens, under non-inflammatory conditions. CD4+CD25– T cells also readily upregulate these markers after TCR activation, but express them only transiently [5,6,7]. However, the use of CD25, CTLA4 or GITR as Treg markers in the presence of inflammation may result in uninterpretable data.
IPEX (immune dysregulation, polyendocrinopathy, enteropathy, X-linked syndrome) is a rare disease, with heterozygous females being asymptomatic, while affected males present severe diarrhea, lymphocyte activation, pro-inflammatory cytokine overproduction, insulin-dependent diabetes mellitus, thyroiditis and immunodeficiency [8, 9]. Scurfy mice are the counterparts of human IPEX. The causative gene in Scurfy mice was identified in FOXP3 which encodes Scurfin, a new member of the forkhead/winged-helix family of transcription factors. Also human IPEX is associated with loss of function mutations in the FOXP3 gene. FOXP3–/– mice lack CD4+CD25+ T cells. A CD4+CD25+ T cell transfer completely prevents autoimmunity, suggesting a correlation between Treg absence and autoimmune diseases [9, 10]. In mice, only CD4+CD25+ T cells express FOXP3 mRNA, while activated CD4+CD25– T cells or differentiated Th1/Th2 cells do not and FOXP3 transduction correlates with CD25, CTLA4, GITR and CD103 expression [10]. However, few data about the role of human FOXP3 are available, and they show some discrepancies with murine models. In humans, there are individual differences in FOXP3 expression. Furthermore, anti-CD3/anti-CD28 stimulation induces FOXP3 expression in CD4+CD25– T cells. These cells may represent the induced Tregs [10].
Tregs develop within the thymus as a functionally mature T cell subpopulation. CD4+CD25+ thymocytes express FOXP3 and present a surface phenotype similar to mature Tregs. In vitro,these cells are naturally anergic to TCR stimulation and exhibit suppressive activity [11].
Thymic development of Tregs requires an interaction between their TCRs and self-peptide/major histocompatibility complexes (MHC), expressed on thymic stromal cells. Studies with double transgenic mice demonstrated that Tregs development requires a TCR with a relatively high avidity for self-peptide/MHCs. However, TCR affinity must not be so high as to lead to deletion. In double-transgenic mice (which express moderate levels of transgenic peptides on thymic stromal cells) most of the thymocytes with a transgenic specific TCR, differentiated into CD4+CD25+ T cells [12, 13]. Treg production was abrogated in the presence either of low affinity transgenic TCRs or a high peptide concentration on thymic stromal cells, probably because of insufficient positive selection or strong negative selection, respectively. Why the relatively autoreactive Tregs are not negatively selected is unclear. Perhaps the superficial co-stimulatory molecules increase the interaction affinity between developing Tregs and stromal cells, as suggested by Treg reduction in CD28- and B7-deficient mice or in CTLA4 immunoglobulin-treated mice [13, 14].
Tregs show a polyclonal TCR repertoire, so they potentially recognize a wide array of antigens. Treg TCRs show a higher avidity for peptide/MHC complexes compared with other specific CD4+CD25– T cells. Because of these features, Tregs may also modulate the responses to infectious agents and downregulate exuberant immune processes [14, 15].
Tregs develop as a functionally mature T cell subpopulation, highly differentiated and able to exert their suppressive activity. When they are stimulated in vitro with a specific antigen, in the presence of CD4+CD25– T cells and antigen presenting cells (APCs), they inhibit effector T cell proliferation in a dose-dependent way [14,15,16]. In vitro Tregs are anergic, so they do not proliferate upon TCR stimulation. However, they require activation via TCR to express their suppressive functions, and this involves both CD4+ and CD8+ T cells [16]. Treg suppression is antigen nonspecific and does not require either TCR re-engagement or the presence of APCs [15, 16]. This may explain the mechanism of linked suppression in transplantation and evidence that histocompatibility is not necessary for Treg function [17].
Treg anergic status in vitro seems closely related to their suppressive function. Indeed, TCR stimulation with high levels of IL-2 or anti-CD28 antibodies abrogates the suppressive activity of Tregs. Removal of IL-2 or anti-CD28 antibodies restores the original anergic state and suppressive activity [18]. Nevertheless, Tregs proliferate in vivo in response to immunization, with no abrogation of their suppressive activity [19, 20]. Treg proliferation seems quite different compared with CD4+CD25– T cell responses, since it is characterized by a defective induction of IL-2, effector cytokines and CD40L. Recent studies showed a reduction in CD4+CD25– T cell number after interaction with a specific transgenic antigen, probably due to the local induction of apoptosis. Diversely, Tregs markedly increased after the encounter with the same antigen, suggesting a reduced susceptibility to apoptosis or an increased recruitment from the circulation. So interactions with autoantigens may shape Tregs repertoire at different sites [19, 20].
Treg suppression may be mediated by soluble factors or by direct cell-to-cell interaction. Treg suppression in vitro seems to correlate with a direct cell-to-cell contact. Indeed, supernatant obtained from Tregs co-cultured with CD4+CD25– T cells does not show suppressive activity, and when Tregs and effector T cells are separated by a semi-permeable membrane, the former fail to suppress the response of the latter [21, 22]. A suppression mediated by cell-to-cell contact may involve competition for APCs (fig. 1). However, APC perturbation does not seem essential as Tregs suppress the effector T cell even without these cells [23].
The role of cytokines in Treg suppression is largely controversial despite intense interest. In vitro neither IL-10 nor TGF-β seem involved as their removal or neutralization do not abrogate suppression. In vivo the mechanisms are more complicated and cytokines are potentially implied in suppressive activity, at least under certain conditions. Indeed, Tregs from IL-10–/– mice show a full capacity for suppressing autoimmune gastritis, but they are not able to abrogate inflammatory bowel diseases in the same mouse. The contribution of IL-10 to Treg suppression is also demonstrated in models of transplantation tolerance, graft-versus-host disease and type 1 diabetes mellitus (T1D) [24]. A recent study by Belkaid et al. [25] showed that during the chronic phase of Leishmania infection, IL-10 production by CD4+CD25+ Tregs is essential for protection against self-destructive inflammation, whereas during the acute phase of disease the contribution of IL-10 is not necessary. The contribution of TGF-β is quite more controversial, although the administration of anti-TGF-β receptor-blocking monoclonal antibody in mice, in the presence of both CD4+CD25– and Tregs, results in neutralization of Treg suppression and in inflammatory bowel disease development [26]. Several other factors are involved in Treg function (IL-2, CTLA4, CD28), but their role in suppression appears controversial [27,28,29].
Finally, Tregs may need cell-to-cell contact to exert their suppressive function both in vitro and in vivo, but in vivo this may be not sufficient. Another possibility is that additional regulatory T cell subsets exert their suppressive function by secreting cytokines, such as the induced CD4+CD25+ regulatory T cells.
CD4+CD25+ T cells can develop from mature, peripheral CD4+CD25– T cells, under certain conditions of antigen stimulation and in the presence of immunosuppressive cytokines, such as IL-10 and TGF-β, vitamin D3, dexamethasone and with CD40-CD40L blockade or immature APCs [30]. Studies in NOD mice demonstrated that iTregs can be produced in vivo, and stressed that their function is generally mediated by immunosuppressive cytokines [31]. Natural Tregs may facilitate the induction of adaptive Tregs from CD4+CD25– T cells. Probably, the two Treg subsets exert their suppressive function in a different manner, depending on the features of the microenvironment, the nature of inflammatory response and the characteristics of antigen stimulation. It is possible that natural Tregs play a dominant role in controlling immune responses under basal conditions, i.e. in a non-inflammatory setting, where a fine balance of the homeostatic process is required and the cell-to-cell interaction seems the most efficient mechanism. On the contrary, induced Tregs probably grant a higher control of the immune response in inflammation settings, where the cell-to-cell interaction is insufficient and amplification of suppressive function may be obtained by cytokine secretion [3, 30, 31] (fig. 2).
NOD mice spontaneously develop diabetes, as result of T cell-mediated destruction of pancreatic β cells. Thus, they represent a useful model for understanding autoimmune diabetes pathogenesis. CD4+CD25+ T cells have recently been described as potentially critical factors in preventing diabetes development. Thus, islet destruction might be the result of an alteration in peripheral mechanisms that normally control and suppress effector functions of autoreactive T cells [32].
CD4+CD25+ T cells are present in low numbers in NOD mice, but they retain a certain capacity to suppress immune responses as demonstrated by the observation that their depletion determines a more rapid disease progression. A recent study demonstrated that Foxp3+ TGFβ+ CD4+CD25+ T cells in pancreatic lymph nodes and islets declined progressively in a age-related way in female NOD mice, whereas male NOD mice were not affected. This decline seems to be chronologically related to the development of severe insulitis in female NOD mice [33].
NOR mice show a similar genetic background to NOD mice, but they do not spontaneously develop autoimmune diabetes. Both murine models are characterized by defects in thymic selection, allowing autoreactive T cell escape. In addition, both NOD and NOR mice spontaneously develop priming of autoreactive T cells, as demonstrated by the observation that their diabetes incidence increases in a pathogen-free environment in the absence of phenomena of cross-reactions. Transfer of CD4+CD25+ T cell-depleted spleen cells from NOR mice to NOD SCID mice induces severe insulitis and the development of diabetes. So the state of protection from diabetes characterizing NOR mice seems to be correlated with CD4+CD25+ T cell function. Specifically, Ott et al. [34] postulated that NOR mice have a higher number of Tregs which are specific for islet cell antigens compared with NOD mice. Several data suggest that organ-specific CD4+CD25+ T cells are highly efficient in abrogating CD8-mediated autoimmune diabetes. So it is possible that a decline only in a Treg subpopulation, specific for islet cell antigens, might be the critical factor that leads to diabetes development in NOD mice [34].
This is consistent with the observation that the transfer in a lymphopenic host of Foxp3-transduced T cells with specificity for islet cell antigens suppresses diabetes progression, while the injection of polyclonal Foxp3-transduced T cells does not have the same result. Thus, Treg homing and activation in pancreatic lymph nodes seems to represent a critical factor for in vivo CD4+CD25+ T cell function [27, 28].
The comprehension of role of CD4+CD25+ T cells in diabetes pathogenesis also has important consequences in the therapeutic field. Bluestone et al. [29] observed that Tregs, derived from BDC2.5 TCR transgenic mice and expanded in vitro, were able to suppress diabetes in vivo in multiple model systems and restored normoglycemia in new-onset diabetic NOD mice. The observation that CD4+CD25+ T cells may have a protective effect on autoimmunity, also after the onset of the disease, leads to considering the possibility of therapeutic cellular vaccines for organ-specific autoimmunity, such as T1D. Several efforts have to be made to improve knowledge about techniques to identify and selectively expand autoantigen-specific Tregs in humans. Once expanded, CD4+CD25+ T cells may be transferred into affected patients to reverse the disease and promote long-term tolerance [29].
Dendritic cells (DCs) are the most potent and efficient professional APCs, thanks to their capacity to detect inflammation signals and induce pathogen-specific responses. Nevertheless, DCs also contribute to maintain tolerance to autoantigens [35, 36].
In order to act as APCs, DCs require a maturation process. DC maturation begins in the peripheral tissues with the activation of toll-like receptors by the antigen, which is consequently internalized. Then DCs migrate through afferent lymphatics into draining lymph nodes, where they present the processed peptides in the context of MHC molecules to naive T cells [37].
However, T cell activation needs additional factors to overcome Treg suppression. IL-6 secretion, induced by toll-like receptor stimulation, may reduce responsiveness of effector T cells to Treg inhibition, enabling their activation [3, 37]. Oldenhove et al. [38] demonstrated that Tregs may exert a certain suppressive effect on mature DCs in vivo. In fact in their study, after elimination of Tregs, these cells stimulated larger Th1 and CTL responses.
Immature DCs (iDCs) are highly specialized in endocytosis while expressing low levels of co-stimulatory and MHC molecules. In the steady state, iDCs may capture autoantigens and present them to T cells in lymphoid organs, with the induction of peripheral tolerance, through abortive expansion, deletion and anergy of the specific T cells [39]. Diversely, if antigen is targeted to iDCs together with anti-CD40 antibodies, promoting APC maturation, T cell activation occurs. In murine intestinal epithelium, the presence of iDCs was demonstrated in the steady state loaded with apoptotic epithelial cells, suggesting that iDCs, located in peripheral tissues, may capture inhaled, ingested proteins or dying cells generated during normal cell turnover. Upon antigen endocytosis, in the absence of inflammation, DCs remain immature and migrate to regional lymph nodes, where they induce peripheral tolerance [40] (fig. 3).
Actually, DC migration and antigen presentation imply at least a partial maturation, with upregulation of MHC and other co-stimulatory molecules. Pulmonary DCs triggered with antigen induce peripheral tolerance and present features of mature DCs [41]. Semi-mDCs may express relatively high levels of MHC molecules but secrete low levels of pro-inflammatory cytokines and present low levels of CD86 and CD40 [37]. TNF-α plays an important role in inducing partial activation of DCs[ 42]. In vitro TNF-α exposed self-peptide loaded DCs were transferred into mice and prevented experimental autoimmune encephalomyelitis [42].
The antigen presentation by iDCs in draining lymph nodes, in the absence of activation signals, may induce Treg differentiation. Dhodapkar et al. [43] demonstrated that injection of mature DCs, pulsed with MHC class I-restricted influenza matrix peptides, determined the expansion of peptide-specific CD8+ T cells, secreting IFN-γ. By contrast, the injection of peptide-charged iDCs increased influenza-specific IL-10-secreting T cells which suppress IFN-γ-secreting T cells. The suppression was contact mediated and IL-10 independent, like Tregs [44]. However, additional factors are probably needed in order to induce Treg generation.
1,25(OH)2D3 represents an important growth factor with a relevant role in several immune processes, thanks to its interaction with the vitamin D receptor (VDR). VDR (belonging to the superfamily of nuclear hormone receptors) is a transcription factor, present in many immune cells where it influences the expression of specific DNA sequences (vitamin D responsive elements). The interaction between VDR and its ligands inhibits DC maturation by reducing IL-12 production and the expression of co-stimulatory molecules, such as CD80, CD86, CD40. Furthermore, VDR activation upregulates the inhibitory receptor ILT3, correlated with DC tolerogenic properties [45].
DCs might play a critical role in T1D development because of their ability to present antigens to effector T cells but also to mediate peripheral tolerance. This is consistent with the observation that the first phase of T1D progression is characterized by the infiltration of the islets by DC, macrophages and B cells [46].
Several studies have documented a reduced DC generation in NOD mice, with bone marrow precursors that preferentially differentiate into macrophages or macrophage-like DC. In addition, the few DCs present in NOD mice seem to be defective in interacting with allogenic T cells, and this may contribute to alter central and peripheral tolerance mechanisms [46].
Angelini et al. [47] demonstrated that DC in children affected by T1D display a lower expression of co-stimulatory molecules, such as B7–1 and B7–2, with a consequent potential alteration in T stimulatory function. This impaired DC phenotype might affect also CD4+CD25+ T cell generation and homeostasis, and enhance autoimmune phenomena [47].
In addition, alterations in DC development have been shown both in children with T1D and in healthy children with a genetic predisposition to develop T1D, suggesting a correlation to the genetic background, independent of disease development [48].
In this scenario DCs may also represent useful therapeutic tools. Tolerogenic DCs, induced by short treatment with 1,25(OH)2D3 or its analogs, probably mediate the property of this hormone to enhance Tregs involved in transplantation tolerance. Furthermore, before insulitis onset, treatment of NOD mice with 1,25(OH)2D3 or its analogs seems to inhibit T1D progression. The slowing of T1D is associated with an increase in Tregs in pancreatic lymph nodes, which may be DC mediated. However, a combination of 1,25(OH)2D3 and steroids in vitro is able to induce differentiation of naive CD4+ T cells in Tregs, even in the absence of DCs [45].
The pathogenesis of type 1 diabetes is T cell-dependent, however, B cells probably play a significant role, even if the exact mechanism involved is still unknown.
An important function of B cells is their ability to act as APCs. Thanks to their receptor specificity, B cells are able to efficiently capture, process and present autoantigens to diabetogenic T lymphocytes. Moreover, once activated, B cells may enhance antigen processing and express a broad array of different peptides on their surface. This may favor the diversification and amplification of T cell responses, by the recruitment of different clones of autoreactive T cells [49].
Studies with the progeny derived from inter-crosses between NOD mice and µMT–/– mice, which lack B cells, show a reduction and a delay in the onset of insulitis in these mice. This suggests that the absence of B cells may confer a sort of protection from the development of spontaneous diabetes. A similar result is obtained in treatment with anti-IgM antibodies. This tolerance status seems to be correlated with an impaired activation of islet-reactive CD4 T cells in pancreatic lymph nodes, suggesting a key role for B cells as APCs for diabetogenic T cells [50].
Moreover, B cell-deficient NOD mice lack T cell responses against 65-kDa glutamate decarboxylase (GAD65) and heat shock protein 60 (HSP60), two of the most important autoantigens in autoimmune diabetes pathogenesis. Falcone et al. [49] demonstrated that these responses are lacking since the activation of GAD65- and HSP60-specific T cells requires the presence of B cells as unique APC.
However, even if with a lower frequency, diabetes can also develop in B cell-deficient NOD mice, suggesting that B lymphocytes are not the unique mechanism operating. In addition, chimeric models with B cells entirely derived from NOD mice are characterized by a tolerance status, demonstrating that B cells may play a pathological role but their presence is not sufficient to determine the occurrence of diabetes [51]. Other alterations have to be present to break tolerance, such as defects in non-B cell APC subsets. This is a critical point because it implies that therapeutic options depleting the B cell compartment (such as rituximab, which has been proposed for the prevention of diabetes) may confer protection from the disease, but probably further interventions, such as manipulations to restore professional APC subsets, are needed to ensure a stabile tolerance [51].
Another important feature of B cell function is related to autoantibody production. A body of evidence documents the presence of autoantibodies in patients with diabetes, also in the prediabetic phases, but it is still unclear if they play a pathogenetic role or are simply epiphenomena. There are several proteins targeted by the autoimmune response, such as insulin, the GAD65 and the tyrosine phosphatase insulinoma-associated protein-2/islet cell antibody 512. The association between some autoantibody patterns and peculiar HLA haplotypes seems to be highly predictive of the progression to diabetes in relatives of patients affected by the disease [50]. Moreover, Greeley et al. [52] demonstrated that the abrogation of autoantibody transmission from a NOD mother to her progeny seems to protect from the development of spontaneous diabetes.
However, autoantibodies fail to transfer disease in mouse models. In addition, reports document the development of autoimmune diabetes in a patient affected by X-linked agammaglobulinemia, suggesting that neither autoantibodies nor B cell function are critically engaged in the pathogenesis of autoimmune diabetes [53].
T1D is characterized by alterations enabling the generation of autoreactive T cells; however, defects in B cell development may also represent a critical event to initiate autoimmunity.
Receptor editing represents a fundamental event in B cell development [54]. At the stage of pro-B cell, the rearrangement of variable (V), diversity (D), joining (J) genes of heavy chains (HCs) occurs, and leads to pre-B cells (fig. 4). Pre-B cells express a µHC, whose synthesis induces light chain (LC) gene rearrangement. LC rearrangement involves first the ĸ locus and leads to a ĸLC. If this process is effective, a λLC will be synthesized. If no LC is appropriate, the developing B cell dies (fig. 5). If the pre-B cell receptor (BCR) is appropriate, unknown signals suppress the V(D)J recombination, with the allelic exclusion at the HC locus. Appropriate signals also grant pre-B cell proliferation and development progression. Inappropriate or underexpressed pre-BCRs stimulate a secondary V(D)J recombination at the HC locus, involving DJ rearrangement or V segment replacement [55]. So, receptor genes are replaced with the generation of non autoreactive pre-BCR and without cell loss. This mechanism is very useful, since autoreactivity is a very common event in B cells. Wardemann et al. [56] showed that the most part (55–75%) of developing BCRs presents self-reactivity.
An effective first receptor editing induces B cell progression and the expression of surface IgM. At this stage, immature B cells are able to interact with antigens and underlie the processes of positive and negative selection. While negative selection deletes autoreactive B cells, positive selection tests the presence of an appropriate signaling, generated by BCR. This signaling, called 'tonic', seems to be the result of a balance between enhancing and inhibiting regulatory molecules, and is ligand independent [57]. B cells expressing signals that are too high or too low escape positive selection and undergo a secondary LC gene recombination. Positive selection is critical, since interaction between autoantigens and B cells, expressing incompetent autoreactive BCR, may result in B cell selection rather than in apoptosis, promoting autoimmune events [58].
Silveira et al. [59] demonstrated that in NOD mice autoreactive B cells specific for soluble self-antigens are not efficiently deleted both in bone marrow and spleen. This defect of tolerance mechanisms may be due to an imbalance between apoptotic and anti-apoptotic molecules or to a weak signal through the autoreactive BCR upon the encounter with self-antigen (as has been proposed for defects in autoreactive T cell selection in T1D). In NOD mice, self-antigens recognized by specific T cells are likely to be soluble, thus autoreactive B cells might represent the most efficient APC subset in expanding diabetogenic T cells [59].
Negative selection correlates with affinity and the signal threshold of BCR. The interaction of autoreactive BCR with self-antigens leads to a new developmental arrest, with the most autoreactive B cells undergoing LC receptor editing [58, 60].
Recent studies in systemic lupus erythematosus (SLE)-prone models, demonstrated an apparent correlation between autoantibody production and consistent LC receptor editing. These apparent contradictions may be explained with the 're-editing' of mature B cells, which leads to autoreactive B cell generation from previously non autoreactive cells. This acquired autoreactivity may derive from alterations in receptor revision, probably induced by B cells carrying a low affinity BCR. In this context, the re-editing process may play an important role in affinity maturation, but may also induce the appearance of autoreactivity [61, 62].
Autoreactive B cells that escaped central selection and entered the peripheral repertoire may become tolerant. The interaction of their BCR with self-antigens determine a cascade of events that leads to a blockade of the differentiation process (functional silencing). B cells also become unable to activate several pathways correlated to BCR activation (anergy). Anergic autoreactive B cells are more susceptible to apoptosis as they do not efficiently home to lymphoid follicular areas compared to normal B cells, and do not produce actively fundamental survival factors [63].
Several studies demonstrated that in murine models of systemic autoimmune diseases, such as SLE, this mechanism is not fully operating, since autoreactive B cells are not blocked in their development after the encounter with the antigen. This process has been evidenced in SLE-prone strains, where systemic antigens may stimulate at high levels and continuously autoreactive B cells. Organ-specific autoimmune diseases, such as T1D, are characterized by limited phenomena of autoimmunity, directed against specific antigens on target organs. Thus, autoantigens are presented only at low levels and they may condition autoreactive B cell function in a different way. A recent study demonstrated that, in a transgenic model of NOD mice, the maturation process and the expression of co-stimulator molecules on the surface of autoreactive B cells are not affected upon the encounter with self-antigen. Only the B cell capacity to proliferate becomes defective. This new tolerance state for autoreactive B cells, characterized by the dissociation between anergy and developmental arrest, may allow them to interact with diabetogenic T cells and promote diabetes onset. Recent studies have opened new scenarios about the role of B cells in the pathogenesis of T1D, with potentially important effects also on therapeutic strategies [64].
Autoreactive T cells may also exert a protective function, at least in the central nervous system (CNS). Indeed, they seem to protect the injured CNS from damage correlated with self-destructive factors produced during inflammation.
In murine models, Yoles et al. [65] demonstrated that autoreactive T cells, specific for CNS myelin basic protein, enhance recovery after optic nerve crush injury or spinal cord contusion. Moreover, active or passive immunization with CNS myelin-associated antigens or passive transfer of T cells reactive to myelin proteins limit the extent of CNS damage. Thus, T cell-mediated autoimmune responses may represent the physiological response to CNS damage [65].
Prof. Maurizio de Martino, MD
Department of Paediatrics, University of Florence
via Luca Giordano, 13
IT–50132 Florence (Italy)
Tel. +39 0555 662 494, Fax +39 0555 703 80, E-Mail firstname.lastname@example.com
Received: January 17, 2006
Accepted: May 15, 2006
Published online: August 11, 2006
Issue release date: August 2006
Number of Print Pages: 11
Number of Figures: 5
Number of Tables: 0
ISSN: 1663-2818 (Print)
eISSN: 1663-2826 (Online)
For additional information: http://www.karger.com/HRP
Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher or, in the case of photocopying, direct payment of a specified fee to the Copyright Clearance Center.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in government regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.


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BCI Drake Division is an industry leader in customer satisfaction and operational quality. Through our creativity in designing solutions for our customers, we have become a premium supplier of corrugated packaging and displays in our region.
Founded in 1969 and acquired in 2016, BCI's Drake Division is an innovative designer and manufacturer of graphic intensive display and signage solutions for high profile retail environments. Long recognized as a leading supplier to some of the most familiar names in consumer retail products, Drake continues to find innovative solutions for diverse packaging requirements in today's ever changing markets.
Visit BCI's Capabilities page to learn how we are designed to fit your needs.
Gary Ames, General Manager
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Alabama Webmaster Training
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Undergraduate Certificate in Web Development
Liberty University , Online
- Ranked among top Regional Universities in the South by U.S. News and World Report in 2015.
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Kaplan University , Online
- Ranked among the Best Online Bachelor's Programs in 2015 by U.S. News and World Report.
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Virginia College , Huntsville
- Instructors are typically real-world professionals with many years of experience in their career fields.
- Ranked #3 in Best for Vets: Career & Technical Colleges 2014 by Military Times.
- Presents the full tuition cost up front. In most cases, even textbooks are included in the total price.
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Stratford Career Institute , Online
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Whatever your motivation for considering an undergraduate or graduate degree, Florida Tech offers programs online that can meet your needs. Here are a few reasons Florida Tech may be the right choice for you.
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Syracuse University Computer Science , Online
- Syracuse University's Master of Science in Computer Science program is designed to help students discover groundbreaking solutions to complex technology challenges and develop expertise in areas such as security and assurance, artificial intelligence, computer architecture, and advanced programming.
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- Founded in 1901, the school has maintained its reputation as a leader in the field, offering a curriculum built on theory, fundamentals, and application with a worldwide community of over 20,000 engineering and computer science alumni
- Syracuse University's College of Engineering and Computer Science is designated by the National Security Agency and Department of Homeland Security as a Center of Academic Excellence in Information Assurance Education (CAEIAE) since 2001 and as a Center of Academic Excellence in Information Assurance Research (CAE-R) since 2009.
- Designed and taught by Syracuse faculty, the curriculum built on theory, fundamentals, and application.
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Use form CCL1TR to appoint a UK tax representative for Climate Change Levy if your business is not based in the UK.
Use form CCL1TR if you need to appoint a UK tax representative because your business makes supplies subject to UK Climate Change Levy (CCL), but has no business establishment in the UK.
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Related forms and guidance
Climate Change Levy: registration form (CCL1)
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Excise Notice CCL1: a general guide to Climate Change Levy
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Excise Notice CCL1/1: registering for Climate Change Levy
Notice about registering for the main rates of CCL and the carbon price support rates of CCL.


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Kentucky Disability Benefits: Social Security Filing and Insurance Options
Find out about Kentucky's state supplemental payments, plus how to appeal a denial of Social Security disability or SSI and whether you qualify for Medicaid.
For Kentuckians with long-term disabilities, disability benefits are available through the Social Security Administration (SSA) programs or their employer's worker's compensation insurance. In some cases, disabled workers may be able to obtain benefits through group disability insurance.
Social Security Disability Programs
There are two SSA programs available to those seeking disability benefits. The eligibility requirements for these programs differ slightly, but some applicants may, in fact, qualify for both.
- The Social Security Disability Insurance program (SSDI) provides benefits to those who have accrued sufficient credits in the SSA Trust Fund by working approximately five out of the last ten years. (Most people who are eligible for Social Security retirement benefits are eligible for Social Security disability insurance benefits.)
- The Supplemental Security Income program (SSI) provides monthly payments to those who have limited income and assets. These claimants may also qualify for other benefits, such as housing assistance, food stamps, and medical care (Medicaid), among others.
How to Apply for Disability Benefits in the State of Kentucky
After submitting the applications forms to the SSA, either through a Kentucky field office, over a toll-free phone line (800-772-1213), or online at www.ssa.org, Kentucky's Disability Determination Service (DDS), which is managed by the Cabinet for Health and Family Services (CHFS) reviews the application.
Contact Information for Disability Determination Services
Here is the contact information for Kentucky's DDS (there are two locations):
P.O. Box 1061
Louisville, Kentucky 40201
Phone: 1-800-928-3202
P.O. Box 1000
275 East Main Street
Frankfort, Kentucky 40601
Phone: 502-564-8050
Fax: 800-928-8050
DDS also determines eligibility for two Kentucky state programs: KTAP (Kentucky Transitional Assistance Program) and Medicaid. For more information visit the DDS section of the CHFS website.
How Much SSI Will I Receive in Kentucky?
The federal government funds a large portion of the SSI payment (up to $710 a month), but the State of Kentucky supplements the federal SSI payment. The amount of the state supplement depends on the recipient's living situation.
- In a personal care facility, the supplement is $520 a month (couples are treated as individuals).
- In a family care home, the supplement is $172 a month (couples are treated as individuals).
- With a caretaker at home, the supplement is $62 a month for individuals, $115 a month for couples.
Kentucky's Cabinet for Health and Family Services (CHFS), Department of Community Based Services, administers the state SSI supplement. You can visit the CHFS website for more information on the supplement.
Kentucky Workers' Comp
All employers in Kentucky are required to carry worker's compensation insurance for their employees, except for certain agricultural employers. That means that anyone who is disabled due to a work accident or illness should file a claim with their employer for workers' comp benefits first. After that, if they remain disabled for at least a year, they may be able to obtain SSA benefits as well.
Disability Rights
For problems not related to cash benefits for disability, you can contact this state agency, which provides advocacy and assistance to those who have specials needs.
Office for Public Advocacy, P&A Division
100 Fair Oaks Lane, Third Floor
Frankfort, KY 40601
800-372-2988 Voice/TDD (Toll Free)
Getting Legal Help with Kentucky Disability Benefits
Disabled persons often require a disability lawyer or authorized representative to help file their disability claim and then appeal the initial denial (only 25% of Kentucky applicants are approved at the initial application stage). Disability lawyers have the experience to help them obtain the proper medical data needed for a compelling claim and the skills to present that case to the Social Security judge. Visit our Kentucky disability lawyers' page if you decide to hire an attorney to help win your claim.
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STARmeterSEE RANK
Down 16,162 this week
Gene 'Bean' Baxter
Gene 'Bean' Baxter was born on November 14, 1959 in Bingley, West Yorkshire, England as Gene Lee Baxter. He is an actor, known for Project X (2012), Rock Star (2001) and Blink 182: The Urethra Chronicles (1999). He has been married to Donna H. Mendivil since October 27, 1991. See full bio »
Born:in Bingley, West Yorkshire, England, UK
More at IMDbPro » Contact Info: View agent


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Department/Office Directory
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Kaplan Humanities Institute
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Kellogg Undergraduate Certificate Program
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Ford Motor Company Engineering Design Center, 2133 Sheridan Road, Evanston, IL 60208
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Agent Ratings for Bill Nesbitt
(61 reviews) Overall star rating: 4.8
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Brian S.
Condo buyer in Las Vegas
Bill made it very easy to buy a place from Los Angeles for an investment property. Scheduled the inspection for me, did the walk through, and other things that made it easy for me to buy the place and feel good about the process. No pressure to buy a place and showed me around a few times. Great experience;)
Win N.
Home buyer in Las Vegas
He is a perfect agent who I can not express all in my mind.Because I am not native speaking english.My english is very limited.But he guided me to right way.I thank him a lot.I believe that he is a special one.
Kathleen L.
Home buyer in Henderson
It was a pleasure working with Bill. He is trustworthy which I think is the best asset for a realtor. He was never "pushy' (which is my pet peeve!) and was very dependable. It was difficult working from across the ocean, but Bill definitely made it less so. We experienced ups and downs due to the distance when faced with what is happening in the market --but we pulled though. Thank you Bill for all your efforts! :)
Home seller in Las Vegas
Bill did an excellent job of helping me with the sale of my home. He explained everything as we went along and made the whole experience as painless and smooth as I could have hoped for. When problems came up, he dealt with the quickly and got us back on track. I would recommend Bill to anyone with a home for sale.
Zach C.
Home buyer in Las Vegas
Bill was wonderful to work with. Being a first time home buyer, I lots of questions and Bill was always available by phone or email to answer those questions. I would recommend Bill to any of my friends or family members who are in the market for a home.
Dennis M.
Condo buyer in Las Vegas
Bill did an excellent job helping resolve several problems that came up during the sale.
Amy L.
Home buyer in Las Vegas
Through Bill we bought two houses in Vegas. We live in southern california and we can't go to view the properties much of the time. So Bill became our eyes during the time of finding property. Bill will go to the houses, take lots of pictures and send to us before we decide of whether to put an offer. Our second house was accepeted last summer, at the time all the properties were having multiple offers and bidding wars. Bill were always very patient and continently viewing and taking pictures of another house after the offer got rejected each time. After several attempts, one of our offer was accepted. The short sale procedure took nine months. But eventually the bank approved the short sale. We are very happy. Bill is an honest, objective and hard working agent. Thank you Bill.
Previous review from Amy L.
No review submitted
Terry R.
Home buyer in Las Vegas
thorough and professional--very satisfied on all levels--prompt courteous and good communication skills--very knowledgeable
Sam/Carol C.
Condo buyer in Las Vegas
We are very pleased with the service Bill provided us. With our very limited time in Las Vegas, he made himself available to us at our convenience and with little advance notice was able to show us suitable properties, from which we made our choice. He patiently explained all aspects of our purchase and provided us with answers to all our questions. We found Bill to be very professional, responsive and a pleasure to work with. We highly recommend him to others.
Michelle W.
Home buyer in Las Vegas
I had moved to Las Vegas in May 2012 and had the good fortune to find Bill Nesbitt through Zip Realty to aid me in my search for a new home. As an Iraq War Veteran and a single mother, Bill was entirely sensitive to my circumstances and was extremely patient in the VA home loan process, which can be quite cumbersome overall. Bill understood how important it was for me to secure a safe, comfortable home for my daughter and I and made the process far easier to handle through compassion, insight into the current market, and informing me on what to expect over the course of time it took to close. I would highly recommend Bill to any close friends or family members as I have the utmost confidence in his abilities as evidenced by his dedication, attention to detail, effective communication, trustworthiness, and intelligence. As a first-time home buyer, Bill made my experience as painless as possible (especially while working with a VA loan) and I am definitely recommending him to any potential buyer, especially my fellow veterans. Bill is amazing and anyone who gets the opportunity to work with him will undoubtedly see wonderful results.
Doug S.
Home buyer in Las Vegas
Living in California and trying to buy a home in Las Vegas was a challenge but after one trip to Vegas and looking at several homes Bill became our trusted eyes knowing our likes and dislikes and determining if we would like the home we were inquiring about or not. The home we finally bought had never been seen by us other than through pictures from Bill until after the offer had been accepted by the sellers and the escrow had started. Bill is without a doubt, one of a kind. You can trust him to do what's right for you. A true man of integrity!! Thanks again Bill for all your hard work.
Debi J.
Bill seemed like a very nice person to work with at first, but doesn't seem interested in helping people see what is available to them. Seems to just want the easy sell.
Response from Bill
Debi was upset because I asked her to get a pre-qualification letter from her lender to make sure she was able to buy a home.
Mike .
worst broker I evere dealt with, never replies to ANY inquires, requested new agent to no avail......................
Response from Bill
I have never ignored a client. I tried repeatedly to reach Mike, but never could.
Sandi F.
My husband and I are very demanding clients and Mr. Nesbitt has been very patient as well as responsive to all our needs. As soon as we saw properties that were to our liking, he immediately acted upon it. A dedicated diligent agent who went above and beyond call of duty. We highly recommend Mr. Nesbitt and we consider him our agent in Las Vegas area henceforth.
Patty S.
Home buyer in Las Vegas
Bill was very patient and informative, I felt really taken care of through the whole home buying process and I feel that he is very trustworthy. Would definitely recommend him to anyone needing a real estate agent.
Gary J.
Home buyer in Las Vegas
He was very helpful in guiding us through the Las Vegas home market
Home buyer in Las Vegas
Bill surpassed his responsibilities as a real estate agent. His knowledge and honesty about different areas of Las Vegas was extremely helpful. We would not hesitate to recommend him to other potential investors.
Craig B.
Home buyer in Las Vegas
Bill is awesome! Living on the East Coast makes it a little uneasy, but Bill made it feel like we were next door. I relied on him being my eyes and ears and truly felt he had my best interests at heart. I would work with him again in an instant!
Home buyer in Las Vegas
Since we were moving from another state, we asked Bill to take pictures of houses that we were considering. He usually went out the same day and had the pictures to us in a matter of hours. He really helped us narrow our search until we found just the right house in our desired neighborhood.
Giau H.
Home buyer in N Las Vegas
Bill is an excellent agent and very professional. In addition, he is very punctual and helpful. We're very glad to have as an agent.
Home buyer in Las Vegas
Bill conducts himself courteously and professionally from beginning to the time I received my home keys. He is also very informative throughout the purchasing process.
Home buyer in Las Vegas
Very professional and looked out for your interest in the transaction.
Mr. Nesbitt is very professional and knows his field well. He goes the extra mile to help.
James S.
Home buyer in Las Vegas
Bill was great through out the process. When we first started looking we really had no idea what we were even looking for. Bill patiently worked with us through the entire process until we eventually narrowed things down to the type of home and location we wanted. Great comunicator!!!!
Jeannie M.
Home buyer in North Las Vegas
He was the best and I have already recommened him to others.
Home buyer in Las Vegas
No review submitted
Initially I was assigned another agent w/ ziprealty. I was not getting good service from him, but he was able realize that and offer to change the agent to Bill. Ideally, it would have been nice to get someone assigned who is not overloaded from day 1. Bill has been great in responding to my requests.
John S.
Home buyer in Las Vegas
Excellent agent who stays on top of every detail of the purchase process and returns phone calls and answers questions in an extremely timely manner. I would highly recommend Bill and plan on using him for my next Las Vegas purchase.
Gordon B.
Condo buyer in Las Vegas
Bill proved to be an excellent realtor. He was always willing to go the extra mile for his customers.
Severino C.
Home buyer in Las Vegas
We bought a vacation house in Las Vegas, Nevada, in which Bill Nesbitt was our realtor. He was patient enough to conduct a long-distance transaction because my wife and I live in Washington state. Every time we were interested in a home that we saw on the Zip Realty website, he became our eyes and went to the houses to inspect and honestly report back to us the pros and cons of each house. He really looked after our best interests, unlike many other realtors that are after making a quick sale and getting a commission. We will definitely refer him to any of our friends that are interested in buying a home in Las Vegas.
Antoniette G.
Home buyer in Las Vegas
Bill is a diamond in the ruff and Zip should be honored to have him as an agent. Bill took care of all of my needs, concerns, issues, and the myriad of details that come up during a house search and closing. I can be very demanding but Bill was always professional, courteous, respectful and concerned. THANKS BILL! I was out-of-state and had to rely on him to set up everything: from researching homes, taking photos of homes, continual communication, picking me up and dropping me off at hotels, advising me on the most appropriate homes, setting up appointments, guiding me through the contract signing and it didn???t stop there. Bill conducted the walk through and made certain everything was in order. I was really impressed by how he was able to navigate through the problems that naturally come up during the process and Bill always came out dignified and I satisfied. I could go further but I think the sentiment is communicated. Any homebuyer would be very fortunate to have Bill as their agent and I greatly appreciate his hard work. It is with deepest gratitude and complete confidence that I highly recommend Bill Nesbitt.
Karen M.
Home buyer in Las Vegas
Bill went beyond my expectations! I will be using him in the future.
Home buyer in Las Vegas
Bill is a true professional. He partnered with me in identifying potential properties and helped me all the way with closing the transaction. I was doing the purchase remotely (being in California) and Bill served as my agent on the ground.
Bill was GREAT. He was always available and provided excellent advice.
Home buyer in Las Vegas
Bill was an excellent Realtor. He was very helpful and was always quick with getting back to me, whether it was answering a question, or scheduling a visit to a prospective house. He was always very knowledgeable and professional. I highly recommend Bill Nesbitt.
Kathy L.
Condo buyer in Henderson
Bill was not only excellent in keeping me updated on the properties available to me, he also was as versatile as I was when my needs changed and they changed so many times from my initial contact that I was surprised he didn't tell us to contact him when we were sure of what we wanted. We went from multiple property types, to maximum price to new areas and if this were a dance; Bill stayed in perfect step with us each step of the way. This was a long distance purchase and Bill stayed on point with us advising where we were and what we needed to do. He advised us of how long we would possibly wait from submission to response which allowed us to keep our stress levels low because of the clear expectations already set by Bill. This was truly a wonderfully stressless experience thanks to BILL.
Townhome buyer in North Las Vegas
bill was so good to talk to not that over powering type and in plain langue so down to earth , it was apleasure to deal with /con crowley
Ryan I.
Home buyer in Las Vegas
Bill is the greatest agent I have ever worked with. As a first time buyer the process was at times overwhelming but Bill was always there for me. He is a true professional and a good friend. I would perosonally recommend him to every one I know.
Townhome buyer in Las Vegas
Bill Nesbitt was very patient, knowledgable and extremely helpful. He was a great pleasure to work with.
Vamsi N.
Home buyer in Las Vegas
If you are a first time buyer or an investor, your answer for right agent is Bill. He has immense knowledge on Las Vegas real estate market and constructions. He is very honest, very punctual and drives your home search in a positive direction. He gives patient ear to your issues and concerns all through the process. If it is not Bill, I could never get the beautiful home I purchased recently. I will take another property for investment with him soon. Keep smiling Bill. You are the best.
Marie I.
Condo buyer in Las Vegas
Bill was superb in every way. He answered numerous questions for me on short sales, title, escrow, inspections, home warranty, closing, HOA, and more. He not only did this by email but even made long distance phone calls and answered each call I made to him to be sure I understood everything. He was kind and patient with us and did not rush his answers but went into detail to be sure we understood everything! We want to purchase more real estate in Las Vegas and want to deal with Bill because he is a true professional!
Mike D.
Home buyer in Las Vegas
Provided great insight into the process. Plus he always answered my phone call
Shaun T.
Home buyer in North Las Vegas
Bill was my first Realtor, and he made the experience wonderful. He was kind enough to teach me the ins and outs of home buying, and keep me aware of things to watch out for, and always in the loop on the process. He did great, and I would recommend him to anyone!
Sylvia D.
Home buyer in North Las Vegas
Very knowledgeable, helpful, and professional. Will definately refer him to friends and family.
Home buyer in Las Vegas
Bill was most helpful to us because of his experience in the investment property area. He was responsive to our questions. He's still providing follow-up. We really liked working with him. He's personable and knowledgeable. Our only area of concern is that when we made our initial offer there was an area that said we had to pay a $100/day penalty if our lender missed the proposed closing date. We questioned this item, because we've bought houses before and have never had to sign something like this. He said it was typical nowadays and that there shouldn't be any problem meeting that deadline. Well, Bank of America missed it and we were charged $700 in late fees. I really wish I hadn't signed that offer, and had instead listened to that little voice that said it just didn't sound right. But I signed it and have paid the penalty. We're thinking of buying another home for rental, and rest assured, we will not sign anything with that clause in it.


text: 1/2 Black Heron are one of many bird species you may spot during this experience. Photo: GERRIT DE VRIES / SHUTTERSTOCK Photo 2/2 Cape Verde is one of the best places to find the red-billed tropicbird. Photo: DAVID THYBERG / SHUTTERSTOCK Photo Previous Next View related excursions Birdwatching in Santiago Get a birds eye perspective on Cape Verde! An experienced guide, trained by an Austrian geographer and nature photographer, will take you to see countless species of wondrous birds. Last edited 21/12/2015 Rare bird species Explore the nature Birdwatching with trained guides Notices Not suitable for wheelchairs Difficulty level Suitable for most Duration Approx. 7.5 hour £ 108 This unique bird watching experience offers the chance to see all the birds you will wish to see on Santiago Island, and perhaps a few more besides, as the guides know places where rare vagrants may show up. Of course the endemics are the highlights: Cape Verde Buzzard (Buteso bannermanni), Cape Verde Warbler (Acrocephalos brevipennis), Alexander's Swift (Apus alexandri) and Jago Sparrow (passer iagoensis). Special desert birds include Cream-coloured Courser (Cursorius exsuls), Hoopoe Lark (Alaemon alaudipes) and Black-crowned Sparrowlark (Eremopterix nigriceps). At the reservoir in Ribeira Secca there might be up to 10 different herons: the endemic Purple Heron (Ardea purpurea bournei), plus Grey, Great White, Little White, Cattle Egret, Squacco, Night, Western Reef. Black Heron (Hydranassa ardesiaca) and Intermediate Heron (Egretta intermedia) are vagrants from Western Africa. Waders include stilts, plovers, sandpipers (sometimes an American spotted one!), snipes, whimbrels, and turnstones. If you have time enough wait until nightfall: the Barn Owl and the rare Little Shearwater (Puffinus boydi) might reward you! The guide will pick you up at the hotel or ship and will care for drinks and food. Austrian geographer and nature photographer, Dr. Herbert Bödendorfer has trained the guides so you are in good hands. The vehicle will be an open Hilux, which offers good visibility. Wear good shoes, a sun-hat and take a windbreaker and or shawl. Language: English Walking information: Hiking footwear recommended Print Share this page Facebook Twitter Google+


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Build-A-Bear is on the hunt for a new e-commerce platform
CEO Sharon Price John says Build-A-Bear's old e-commerce system is a big reason for disappointing online sales in December.
Build-A-Bear is on the hunt for a new e-commerce platform
CEO Sharon Price John says Build-A-Bear's old e-commerce system is a big reason for disappointing online sales in December.
What is Wal-Mart's e-commerce strategy?
Wal-Mart's acquisition of Moosejaw is the latest in a series of recent moves that illustrate the retail giant's lack of a coherent e-commerce strategy, ...
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A total of four execs leave in a shakeup, and two new board members will join as Twitter works on a turnaround.
Posted 01/25/2016Bloomberg NewsPost a comment
Twitter made its interim leader permanent, and Dorsey named Adam Bain Twitter's chief operating officer.
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Dick Costolo stepped down as Twitter CEO on June 11 and no replacement has emerged. But the social media platform has remained busy in the interim.
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While the social network's ad revenue grew 63% year over year in the second quarter, it only added 8 million new users. Interim CEO Jack Dorsey says that isn't good enough.
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Mobile advertising accounted for 89% of its overall ad revenue. The social network also bought marketing vendor TellApart.
Posted 04/29/2015Zak StamborPost a comment
Twitter added only 4 million new users in the fourth quarter. But CEO Dick Costolo says the social network is already turning that around in the current quarter, and the company is projecting at least 4.5% growth this quarter in consumers using the social network.
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The two companies had a similar deal that expired in 2011.
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500 million a month, in fact. And Twitter's latest plan to lure more of those lurkers to sign up for the social network is to make Twitter easier to use. A new feature will remove the need for people and brands to follow accounts when they first join the social network.
Posted 11/12/2014Zak StamborPost a comment
Madbits focuses on developing technology to understand and organize information about images. The move comes as the social network is increasingly targeting visual content.
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Shoppers also can access TV shows from the microblogging service.
Posted 10/10/2013Zak StamborPost a comment


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Becoming an Outdoors-Woman Winter Workshops in North Dakota
The North Dakota Game and Fish Department's Becoming an Outdoors-Woman program has scheduled one-day winter workshops Jan. 26-27 at Cross Ranch State Park, Center, and Feb. 24, 25 and 26 at Lake Metigoshe State Park, Bottineau.
Cross-country skiing (Jan. 26) and snowshoeing (Jan. 27) will be offered at Cross Ranch.
Lake Metigoshe offers a variety of classes. A class on dog sledding for first-timers only is available all three days with a limit of six participants per session (this class fills quickly). Snowshoeing and tracking, winter survival and cross-country skiing are available Feb. 25 and 26.
Women interested in the workshops are encouraged to register online at the Game and Fish website, gf.nd.gov. Those interested should sign up immediately, as each class is limited to 12 participants. The cost is $75 for dog sledding and $50 for the other classes. Sign up for a class on two days and receive a $15 discount. Preregistration with payment is required. Equipment and snacks will be provided.
An information and registration form is available for download by accessing the department's education/outreach link, or by contacting Nancy Boldt, North Dakota Game and Fish Department, 100 N. Bismarck Expressway, Bismarck, ND 58501-5095; (701) 328-6312; or email firstname.lastname@example.com.
Sounds like a great program
Submitted by Retired2hunt on Tue, 2011-11-29 16:22.
Sounds like a great program that North Dakota offers to women wanting to become more involved or educated on the outdoors and outdoor activities. A great focus to get more females into the outdoors and hopefully into the sport of hunting. There are definitely two workshops within the program that would be of great benefit to the participants and this is the tracking workshop as well as the outdoor survival workshop. The dog sledding sounds pretty cool to me as well. Anyhow a great program with some excellent workshops. Kudos to North Dakota with their efforts on generating more woman involvement.


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