Total cases of MSDs developed by workers in Mexico reported by the IMSS [8, 9].
\\n\\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 191 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 261 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
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The human muscle strength measurement is important for understanding human capabilities. Nevertheless, the knowledge about strengths developed by an individual during work does not give the specialist enough information to solve ergonomic problems that lead to musculoskeletal disorders (MSDs). Thus, a work-system elements assessment should be necessary to find hazards that cause microtraumas [1]. The microtraumas outrun the body’s recovery system causing work-related injuries that result in musculoskeletal disorders (MSDs).
\nFrom a mechanical point of view, when a machine repeats intensely specific movements during its operation, the applied forces cause fatigue in its mechanisms [2]. From a biomechanical point of view, the musculoskeletal system suffers from fatigue and wears down in joints and muscle injuries [3], when there are ergonomic risk factors at work such as employees’ prolonged exposure time to awkward postures, excessive force exertion, repetitive movements, and manual material handling, causing fatigue and impacting on the health and well-being of workers [4, 5].
\nConsequently, a correct identification of ergonomic risks and their physical location in production areas becomes vital for the prevention of work-related illnesses.
\nIn Mexico, MSDs were included in the work-related illness classification by the Mexican Health Secretary (SSA) until 2008. The Mexican Institute of Social Safety (known as IMSS) categorized the information about MSDs into seven diseases and one injury. This catalog is named “MSDs classification according to a kind of injury” and contains the following diseases [8]:
Other synovitis, tenosynovitis, and bursitis
Radial styloid tenosynovitis (Quervain)
Shoulder injury
Carpal tunnel syndrome
Epicondylitis
Other enthesopathies
Osteoarthritis/arthrosis
Dorsopathies
In 2009, the number of MSDs was recounted for the first time. Table 1 presents data of a nine-year period (2009–2017) [6, 7]. During that period, the IMSS reported only 20,523 cases, showing an underreporting problem. Aspects like authorities not properly reporting risk conditions or workers’ fear of being fired if they notify symptoms, as well as employers’ evasion of mandatory law compliance [8, 9], contribute to the problem of lack of information. Despite work related-illnesses not being appropriately studied as MSDs yet, there was a data ascendant tendency in the results; see Figure 1.
\nYear | \nWork-related illnesses | \nWork-related injuries | \nPercentage of MSDs with respect to work-related injuries | \n
---|---|---|---|
2009 | \n266 | \n4101 | \n6.49 | \n
2010 | \n513 | \n3466 | \n15.80 | \n
2011 | \n788 | \n4105 | \n19.20 | \n
2012 | \n1309 | \n4853 | \n26.97 | \n
2013 | \n1893 | \n6364 | \n29.75 | \n
2014 | \n2604 | \n8301 | \n31.37 | \n
2015 | \n3722 | \n12,009 | \n30.99 | \n
2016 | \n4273 | \n12,622 | \n33.85 | \n
2017 | \n5155 | \n14,159 | \n36.41 | \n
Total | \n20,523 | \n69,980 | \n29.33 | \n
Data ascendant tendency of MSD cases developed by workers in Mexico [
In Mexico City, during the first forum on safety and health at work carried out in August 2015, it was determined that MSDs will be subjects of care because of their impact on workers’ health [10]. To abate this health problem, the Mexican Ministry of Labour and Social Safety Secretaria del Trabajo y Previsión Social (STPS) issued a mandatory rule called Federal Rule for Safety and Health at Work (Reglamento Federal de Seguridad y Salud en el Trabajo) in November 2014 [11]. It includes a new employer obligation for identifying, reporting, and reducing ergonomic risks inside facilities. Now, ergonomic risk factors are highlighted. Hence, the correct identification of MSDs becomes a big problem for the employers. Thus, the identification of ergonomic risk and musculoskeletal disorders (MSDs) for their prevention is too important. The aim of identifying the risks is to find process conditions that lead to musculoskeletal disorders and work-system elements that need changes from an ergonomic point of view.
\nIn this chapter, a methodology for detecting musculoskeletal disorders was employed in a case study, and the diagnosis and analysis developed were used to propose a mapping representation of risks inside the workplace. The mapping resulted in a standardized representation of risks to ensure risk identification. The method includes (a) reports of the employees’ complaints about workstation design and symptomatology suffered by workers as input information, (b) a description about the nonergonomic elements of the task and biomechanical studies on ergonomic risk factors that cause MSDs, (c) risk assessment results and work-related injuries and illnesses as output information, and (d) the cost of nonergonomics spent per year giving extra information that supports the decision-making about future ergonomic interventions and workstation redesign.
\nThere are representations of conditions from an ergonomic point of view, for example, the empirical design of a human-machine system about the person-process relationship designed by Lemon [12], which involves an analysis about job organization, environment, and workplace; however, his model did not consider the work-system inputs/outputs, that is, inputs, information about risks and hazards that lead to MSDs and outputs, and information about the consequences of risk factor exposure. Axelsson’s design [13] included work-systems and quality in a model of ergonomics. The model combined the concept of “fitness for use” developed by Juran [14] and the concepts in the book “Fitting the Task to the Man” developed by Grandjean [15]. His model based on Lemon’s proposal considered only the work-system inputs, like interaction between the worker and the process, inside work space but did not include the outputs. Delgado-Bahena et al. proposed the Ergonomic Hazards Mapping System (EHMS). The model was developed using a rough layout in which the body parts exposed to hazards or risk factors were identified [16]. Nevertheless, their model only considers the work-system outputs and did not provide information on what leads to MSDs. It is important to consider that the models presented above contribute to understanding the ergonomic process problem, but they do not add information for detecting and preventing MSDs.
\nIn the industrial context, a system comprises an interacting component collection that brings together common purposes; the system is limited by variables at any moment in time and is subject to a cause-effect mechanism [17]. The process mapping schematizes the system model using a pictorial relationship between variables. It is composed of legends, symbols, and scales explaining the interactions between system elements with the aim to identify the activities that add value [18]. It is divided into three parts: input-process-output, where the input connections or linkages among a selected part of a process (work system) transform the resources into another valued form (output). The process map represents the whole (end-to-end) work process [19]. Therefore, designs of ergonomic risk map based on the process map concept can contribute to identifying the risk of developing MSDs.
\nThe workplace comprises three polish-screeners designed and built by company personnel, and they were used for polishing pieces of metal. The three machines polished around 50,000 pieces daily. The production time comprised three shifts of 8 h, with three operators per shift. The task was developed on a standing posture. Workers took a lunch time of 0.5 h, at the middle of the work period. Ergonomic risk factors like manual material handling, repetitive movements, awkward postures, and force exerted were to be identified as a part of task performance as is observed in Table 2.
\nWork method.
To design the mapping, an analogy between the process map elements and ergonomic risks was developed. The relationship map regards the input/output connections or linkages among selected work tasks, and workstations were defined. The result is presented in Table 3.
\nProcess components | \nErgonomic risk map | \n
---|---|
Input | \nInformation about risks and hazards that lead to MSDs \n
| \n
Process | \nInformation about how the risk exposure leads to MSDs \n
| \n
Output | \nInformation about the consequences of risk factor exposure \n
| \n
Analogy between the process map elements and the ergonomic risk map elements.
The notations used for classifying the body segment affectation and the risk level of developing MSDs were based on the concept used by the ergonomic standards ISO 11228-3 [20] where the color identification for each risk level was as follows:
Green—there is no risk of developing musculoskeletal disorders; a change in working conditions is unnecessary.
Yellow—there is a risk of developing musculoskeletal disorders; a change in working conditions is needed.
Red—there is a high risk of developing musculoskeletal disorders; a change in working conditions is needed immediately.
Information developed during the work-system assessment that can contribute to identifying risks that cause MSDs was organized according to the connections or linkages between ergonomic risk map elements as follows:
\nInputs: work place conditions and human factors
Work place design
Nonergonomic task content
Individual characteristics
Processes: ergonomic risk factors that cause MSDs and force demands by shift required to perform the task
Weight manipulated
Body segments affected
Color identification of the risk level
Force demands by shift required to perform the task measured in Newtons by movement
Number of repetitions of the exertion strength
Outputs: risk assessment results, work-related illnesses, and the cost of nonergonomics
Pain points in body segments
Resume of task assessments
Work-related injuries and illnesses
Cost of nonergonomics
Step 1. A list was made with risks or hazards that had been identified in a workplace, using data from assessment checklists. It should include only the workstation elements that limit the overall movement of the body or increase force requirements, causing pain or discomfort.
\nStep 2. A list was made with nonergonomic task elements, like awkward postures, repetitive movements, force exertion, and insufficient time recovery, among others.
\nStep 3. A list was made with individual characteristics that workers should change to prevent MSD development.
\nStep 4. Photographs were added to identify the manipulated weight in each task element.
\nStep 5. Workstation layout was added. It represented machinery used for developing tasks.
\nStep 6. Images of body segments affected with color identification of the risk level were added.
\nStep 7. A list was made with force demands that caused pain/discomfort and exceeded the permissible standard value. It included isometric strength, leg lifting strength, grip strength, push and pull (initial force/kept force), and dynamic back extension strength, among others, in Newtons. The analysts were free to choose the measurement method that they consider most appropriate to complete this section.
\nStep 8. The number of repetitive exertions developed by workers was included.
\nStep 9. A drawing of a body segment that identifies a point of pain was added. It represented the pain symptoms suffered by workers.
\nStep 10. All the simple risk assessments developed to determine the acceptability of risk were provided in one table. Identification of results with a color according to each risk level was obtained. REBA, NIOSH equation, and OCRA among others were included.
\nStep 11. All the work-related injuries or illnesses suffered by workers were categorized according to frequency in a Pareto chart.
\nStep 12. The cost of nonergonomics was estimated. It comprised workers with work-related injuries or illnesses, the daily salary (it included allowance for temporary inability and replacement worker salary), an average of lost workdays by a worker, and the total lost workdays per year.
\nThe ergonomic risk map was implemented in three polish-screener machines used for polishing pieces of metal; only nine workers were assigned to develop this task. The machines were poorly designed and built by engineers from the company. The workstation design did not consider basic anthropometric requirements, and this situation caused insufficient space for legs, incorrect working height and inconvenient arm reach, producing awkward postures that cause pain and discomfort.
\nWith respect to nonergonomic task content, the risks found were as follows: exerting excessive force, similar task repetitively, doing work in awkward postures, being in the same posture for a long period, coming into contact with vibration surfaces, and manual handling—pushing and pulling loads and lifting and carrying loads; these conditions caused microtraumas that affect the body’s recovery system of the workers.
\nMoreover, individual characteristics like poor work practices, poor fitness, poor health habits, and poor work readiness add a probability of developing MSDs. Thus, programs about healthy life and better practices of manufacturing should be implemented.
\nThe work method included three task elements with manual handling—lifting and carrying loads:
Barrel filling with metal parts: a filled cardboard with 30 kg of weight is lifted over the shoulder 44 times, exerting an excessive force of around 277 N in each lifting,
sieving the corn cob powder from the metal parts: a filled metallic bucket containing metal polish with 20 kg of weight is handled 2295 times, exerting an excessive force of 77.62 N in each grip strength, and
moving filled cardboard containers with 80 kg of weight to an inspection area 310 times. Leg lifting strength of 143.20 N, dynamic back extension strength of 245.15 N, and push and pull (initial force/kept force) of 291/236 N, respectively, were considered in this force demands.
The task exceeds the biomechanical work load capacity of workers; this means that the musculoskeletal system suffers from fatigue and wears down in joints and muscle injuries. The workers have developed dorsopathies.
\n\n
Upper limbs—Red—there is a high risk of developing musculoskeletal disorders; the repetitive movements need to be eliminated immediately.
Shoulders—Red—there is a high risk of developing musculoskeletal disorders; the height of the barrel need to be reduced immediately.
Trunk (back)—Red—there is a high risk of developing musculoskeletal disorders; the conditions of manual material handling need to be changed immediately.
The method used for the classification and definition of human muscular strength was proposed by Mital and Kumar [21], which divides the strength criteria into two sections: characteristics of the effort that include static isometric muscle strengths and isokinetic muscle strengths and characteristics of the application that include static functional strengths and dynamic functional strengths. The results obtained are summarized in Table 4.
\nClassification | \nMeasurement by movement (N) | \nNo repetitions | \n
---|---|---|
Isometric shoulder strength | \n277.00 | \n44 | \n
Leg lifting strength | \n143.20 | \n310 | \n
Grip strength | \n77.62 | \n2295 | \n
Push and pull (initial force/kept force) | \n291/236 | \n310 | \n
Dynamic back extension strength | \n245.15 | \n310 | \n
Force demands by shift required to perform the task.
In order to determine the pain points in body segments a questionnaire about MSD symptoms was to apply to the 9 operators of the three polish-screener machines. In the questionnaire the workers had to mark the body segment where they felt pain or had any injury. The resume of their answers is shown in Figure 2. The results do not correspond with the official information provided by the safety and health department used for building the Pareto chart developed for determined work-related injuries and illnesses (see Section 5.3.3).
\nPain points in body segments selected by the workers through a questionnaire.
The results from the simple risk assessments were summarized in a table. In all the cases, the resulting risk levels were unacceptable. It allowed identifying the main unsafe and unhealthy task components. See Figure 3.
\nIdentification of task assessments.
The method employed to represent the work-related illnesses was the Pareto chart. It is a frequency distribution (or histogram). It was used for arranging injuries and illnesses by category. The Pareto method and rules of 70/30 (Pareto principle) allow identifying the main MSDs developed by workers in the work area. It can be used from the ergonomic intervention standpoint [22]. The information to build the Pareto chart was proportioned by the safety and health department. This official information indicates that all workers in the area (nine in total) have been suffering from almost two work-related injuries or illnesses (see Figure 4). It confirms the analysis developed in Section 5.2.1. However, it is contradictory with respect to workers’ complaints. They identified the shoulder pain as the main symptom of MSDs. Thus, new studies to implement strategies to balance the differences of opinion are necessary.
\nPareto chart about work-related injuries and illnesses.
Workers with work-related injuries or illnesses: 18.
\nDaily salary: $34.63 USD (includes allowance for temporary inability).
\nAverage of lost workdays by a worker: 35.
\nTotal lost workdays per year: 630.
\nTotal cost of nonergonomics: $392,704 USD ($7,461,380 MXP).
\nThe resulting cost supports the suggestion to change the working method to eliminate repetitive movements, reduce the barrel height, and improve conditions about manual material handling.
\nAll the information presented in previous sections was organized in a single spreadsheet. The ergonomic risk map shown in Figure 5 summarizes result series derived from an exhaustive work place evaluation. The map added evidence necessary to determine that musculoskeletal disorders were caused by the workplace and incorrectly designed tasks. The resulting ergonomic risk map allowed to determine the causes of MSDs developed in activities in a three polish-screener, establishing the barrel height as a main cause of risk. The excessive height forces the material handling above the shoulders. This increases force demands required to perform the task. On the other hand, the work method must be changed in order to reduce repetitive movements. The map improves the employers’ understanding about the origin of ergonomic problems present in the polishing area and supports the decision-making about improvement projects focused on risk elimination.
\nErgonomic risk map from three polish-screener machines.
The assessment and diagnosis method used for building an ergonomic risk map was developed and implemented with the objective of identifying the relationship, between the workplace design and the nonergonomic content task. The standardized method allows obtaining relevant diagnosis about hazards and ergonomic risks factors present in the work system that leads to musculoskeletal disorders. The study shows that the ergonomic risk map (a) improves the understanding of the workers and employers about the origin of ergonomic problems present in working areas, (b) identifies the main unsafe and unhealthy areas and work-system components, (c) supports the decision-making about improvement projects focused on risk elimination. However, the complaints and employers´ opinion in many cases were contradictory with respect to official information. Thus, new studies to implement strategies to balance the differences of opinion are necessary.
\nThe authors are thankful to the company and its workers for their disposition, the Chemistry and Engineering College for its support, and industrial engineering students for their enthusiastic participation.
\nB lymphocytes play vital role in maintaining the normal immunologic functions of the body. Their functions range from producing antibodies to presenting antigens. They are also involved with productions of several regulatory cytokines, such as IL-2, IL-4, IL-6, IL-10, IL-12, TGF-β1, TNF and IFN-γ. Each of these functions are further fine tuned by the fact that they are dependent upon several factors, including the B lymphocyte subsets, their location and the type of stimuli that is encountered by the specific B cell subset in that particular environment. Understanding all these components of B cell development is not only required for getting a better picture of B cell biology, but is also necessary for understanding various immunologic anomalies that lead to disorders. They are also important towards generating effective B cell based therapies. This chapter begins with explanation of the overall pathway of B cell development and the organs that are involved in its various stages. This is followed by discussion on the role of gene rearrangements in the entire process. Subsequently, the role of various transcription factors has been addressed.
\nDuring foetal life, B cells are generated in the foetal liver. Subsequently in the adults, they are produced by differentiation of haematopoietic cells (HSCs) in the bone marrow [1]. Most of the stages of B lymphocyte development take place in this primary lymphoid organ. The pluripotent HSCs gradually differentiate into progenitors, which have increasingly lower potency. Initially, they form a population of cells that are known as multipotent progenitors (MPPs). These progenitors, in turn, give rise to two main progenitor populations: common granulocyte/megakaryocyte/granulocyte progenitor (CFU-GEMM) and early lymphoid progenitor (ELP). CFU-GEMMs subsequently develop into cells that have either myeloid or erythroid potential. On the other hand, cells with lymphoid potential arise from ELPs. Thus, CFU-GEMMs are the primary source of those elements of blood that are non-lymphoid in nature, whereas the lymphoid elements originate from ELPs. Two major precursors arise from the ELPs, common lymphocyte progenitor (CLP) and early T-lineage precursor (ETP). Both Pre-NK cells and Pre-B cells develop from CLPs, which eventually give rise to NK cells and B cells, respectively. T cells derive from thymocytes, which are generated by differentiation of the ETPs. The CLPs give rise to early Pro-B cells first. They mature to form the late Pro-B cells, which eventually develop into Pre-B cells. Immature B cells arise from these Pre-B cells and they leave the bone marrow to enter into the secondary lymphoid organs. Subsequent stages of B cell development primarily continue in the spleen. During this entire period of maturation, the various B cell subpopulations are found to migrate within the bone marrow, keeping in touch with the stromal cells. Initially, the progenitor cells having highest potency lie in the endosteum. This region is located near the inner surface of the bone. As the B cell progenitors mature to give rise to cell types that have less potency and are more committed towards the B cell fate, they start migrating towards the central sinus of the bone marrow cavity. During this entire process, these maturing cell populations are reported to remain in contact with the reticular stromal cells, which are believed to provide indispensable signals for migration and maturation. This process continues till the developing cell reaches the stage of Immature B cell [2, 3, 4].
\nIn addition to HSCs, B lymphocytes (in various stages of development) and plasma cells (PCs), the bone marrow consists of specialised cells that have multiple roles in various stages of this developmental process. Together they form the “niches” that are vital for normal functioning of the different systems associated with the bone marrow. The most important components of these “niches” are stromal cells and regulatory T cells. Out of these, the stromal cells form an extensive network of non-lymphoid connective tissue in the bone marrow [5, 6]. They serve dual functions of forming adhesive contacts with developing lymphocytes and providing cytokines/chemokines/growth factors to them as per requirement. In context of maintenance of plasma cells, earlier reports have indicated that they are provided with survival signals such as CXCL13, IL-6, APRIL and BLγS by the stromal cells [7]. In addition, it has been found recently that the regulatory T cells present in the bone marrow play vital role in maintaining plasma cell pool [8]. The dendritic cells (DCs), present as perivascular clusters in the bone marrow, have also been reported to provide signals that are vital for B lymphocytes [9]. Megakaryocytes [10], eosinophils [11] as well as basophils [12] resident in the bone marrow have also been found to play role in maintaining plasma cell pools.
\nThe immature B cell undergoes final stages of development in the spleen to form mature B cells. The spleen primarily consists of red pulp, white pulp and marginal zone. The red pulp is made up of large, blood-filled sinuses and serves as the blood-filtering system of the spleen. In addition, the splenic macrophages play important role in recycling of iron. The white pulp is organised in line with the lymph nodes and consists of lymphoid sheaths having distinct B-cell and T-cell compartments. The marginal zone is a layer of highly specialised cells that surrounds the white pulp [13]. It plays a very important role in immunity because those haematopoietic cells that remain in circulation (as part of the surveillance mechanism) need to be able to migrate through blood and lymphatic systems continuously. The marginal zone plays a vital role in this process due to its strategic location in the spleen. It has been observed that G-protein linked receptors are involved in the signalling process that is responsible for active transport of B- and T-lymphocytes to-and-from the white pulp [14]. The specialised cells that constitute the marginal zone include two subsets of macrophages, the marginal-zone macrophages and the marginal-zone metallophilic macrophages. The first subset is present as an outer ring and express SIGNR1 (a C-type lectin) [15, 16, 17] and MARCO (a type I scavenger receptor) [18]. The second subset is present as an inner ring, lies closer to the white pulp and expresses SIGLEC-I (an adhesion molecule) [19]. A specialised B-cell population, known as marginal zone B cell, and DCs are located in between these two rings of macrophages [20, 21]. Figure 1 shows the major cell populations that are generated in the bone marrow and peripheral lymphoid organs during the process of B cell development.
\nMajor stages of B cell development.
It has also been reported that the antigens are encountered by the mature B cells in the lymphoid follicles. This process is aided by T cells present in the germinal centres. All the subsequent stages of B cell development, including generation of various Ig isotypes, class switching and somatic hypermutation, contribute towards diversification of the antibody repertoire [22].
\nOn activation by cognate antigen, the activated B cell can either differentiate into antibody-secreting plasma cells/plasma blasts or get recruited into a specialised region known as the germinal centre (GC). Those activated B cells that enter the GC subsequently undergo several rounds of proliferation, class-switching and affinity maturation. Thereafter, the GC B cells that have completed these steps successfully give rise to either long-lived plasma cells or memory B cells [23]. When the B cells undergo these changes in the GC, their transcriptional repertoire also undergoes huge transformation [24]. Figure 2 shows the various stages that are developed after activation of B cells.
\nMajor stages after activation of B cells.
In general, three subsets of B cells derive from naive B cells, B-1 B cells, follicular B cells and marginal zone (MZ) B cells. Furthermore, B-1B cells form two subsets, B-1a and B-1 b B cells [25]. All the subsets can be clearly identified on the basis of their surface markers. These surface markers can also be used to identify their progenitor populations. Table 1 shows the various stages of B cell development and the most prominent cell surface markers that are used to identify them. Each one of these subsets maintains distinct location(s) and function(s). All these subsets of B cells produce functionally important antibodies. However, they vary in huge terms in reference to their origin and function [26, 27, 28].
\nMost prominent cell surface markers used to identify various stages of B cell development.
B-1 B cells have not been successfully studied in mammals, including humans [29]. Thus, most of the findings are based on studies performed in mice. The most interesting finding from these studies is that although the progenitors of B-1 a and B-1 b cells are distinct, they are found to occupy the same locations, namely the pleural and peritoneal cavities. In addition, it has been observed that the environment offered by these cavities plays a significant role in shaping the functional characteristics of these B cell subsets. The milieu of these cavities also influences the functional characteristics of B-2 cells that reside there, although in low numbers [30, 31].
\nSeveral studies have shown that the naive, mature peripheral follicular B cells reside in two main niches during their circulation/recirculation through the bone marrow. Out of these, the “follicular niche” present in the spleen/lymph nodes/Peyer’s patches is the main site that is occupied by these cells. These “follicular sites” are thought to play important role in those immune responses against protein antigens, which are T cell-dependent [25].
\nIn addition, some follicular B cells have also been reported to home in the bone marrow [32]. The site of their homing has been termed as “perisinusoidal niche” and consists of a part of population of the same circulating follicular B cells that are found in the secondary lymphoid organs mentioned earlier. Interestingly, the follicular B cells residing in the bone marrow are involved in T cell-independent immune responses against microbial pathogens harboured by blood [33].
\nMZ B cells mainly home near the marginal sinus of the spleen. This process is facilitated by the molecules SIP 1 and SIP 3, which are receptors for sphingosine-1-phosphate [34, 35, 36]. These cells are mainly involved in T cell-independent immune responses against blood-borne microbes [34]. It has also been reported that these MZ B cells can transport pathogens from the marginal sinus to the splenic follicles, sites where the follicular B cells reside [21, 37]. Moreover, a few
As mentioned earlier, B cell development starts from haematopoietic stem cells (HSCs) in the bone marrow and continues either at the same site or in the peripheral sites. Two cellular pathways are believed to be involved in formation of mature B cells from T2-like cells, one in the bone marrow, and the other in the peripheral sites [25]. Thus, the population of mature B cells present in the bone marrow is heterogeneous in nature. It has been observed that one population of these cells is characterised by sIgMhigh IgDlow CD23−, and do not respond either to BLγS/BAFF or multivalent antigens. In contrast, another population of these cells is sIgMhighIgDhigh CD23+, and responds to both BLγS/BAFF and multivalent antigens [32, 40]. The exact mechanisms that are responsible for this differential developmental pathways and their significance is yet to be understood completely.
\nOnce immature B cells form, they generally migrate to the peripheral sites. These immature B cells have very short half-lives and on engagement with BCRs, they tend to undergo apoptosis instead of proliferation [41, 42]. The immature B cells are also referred to as “transitional B cells” and can be further subdivided into three subsets; T1, T2 and T3 [41, 43, 44]. CD93/AA4 (the B-lineage precursor marker) is expressed in all of these “transitional B cell” subsets. However, they display differential expression of IgM and CD23 on their surfaces that is exploited to identify them. T1 cells are characterised by IgMhigh CD23−, T2 cells by IgMhigh CD23+ and T3 cells by IgMlow CD23− [41]. All data till date suggest that T2-like cells give rise to mature B cells, either in the bone marrow or in the peripheral sites.
\nTolerance to self-reactive antigens can take place by any of the existing three mechanisms: deletion, editing or anergy. In spite of significant number of studies addressing this aspect, it is not yet clear whether this tolerance is achieved by negative selection of self-reactive B cell clones or failure of positive selection. Several studies have suggested that local levels of BAFF may influence this decision [45, 46].
\nIn humans, majority (around 55–75%) of immature B cells have been found to be self-reactive [47], indicating that clonal deletion may serve as one of the main mechanisms of elimination of these self-reactive cell populations. However, no such experimental data is available from mice. In addition, receptor editing has also been found to contribute towards elimination of such self-reactive immature B cells. The process of anergy, although a bit controversial, has emerged as the third mechanism involved in removal of self-reactive B cells [48].
\nSpleen is the main site for positive selection of non-self reactive B cell clones. It has been reported that survival of the peripheral B cell populations is dependent upon continuous signalling through B Cell Receptor (BCR). It holds true for both populations, follicular B cells as well as marginal zone (MZ) B cells [49, 50]. Although a wide variety of V-gene segments are expressed by both follicular and MZ B cell populations, it has been observed that in IgH transgenic mice, those immature B cells that are specific for phosphorylcholine give rise to MZ B cells [51]. In another study, it has been observed that deletion of RAG2 in adult mice results in selective retention of MZ B cells over follicular B cells [52].
\nIn addition to BCR, Notch2 signalling plays a significant role in MZ B cell development [53, 54]. Notch2 is a member of Notch family of receptors and ligands [55]. Interaction between Notch2 and DL1 has been found to be responsible for developing a unique MZB cell niche [54]. Moreover, interaction of Notch2 with NF-κB pathway component p50 helps in maintaining MZ B cell pool, without affecting the pool of follicular B cells [56]. The NF-κB pathway may also work in synergy with the BAFF-BAFF-R pathway, both in context of survival of follicular B cells and generation of MZ B cells [57]. In addition, studies have indicated that BCR signalling and Notch2 signalling may work synergistically for development of MZ B cells [58].
\nThe antibody responses demonstrated by the various subsets of B cells depend upon formation of the fully functional antibody having the required specificity. Each such fully functional antibody is constituted by two light chains (IgL) and two heavy chains (IgH). Each light chain, in turn, consists of variable (V), joining (J) and constant (C) domains. On the other hand, each heavy chain consists of variable (V), diversity (D), joining (J) and constant (C) domains. As mentioned earlier, mature B cells arise by stepwise development from Hematopoietic Stem Cells (HSCs). The genes encoding various domains of the heavy and light chains of the antibody also get progressively rearranged during these developmental phases [59, 60]. Finally, a specific group of genes encoding the various regions of the antibody get expressed [61, 62]. This remarkable process gives rise to the huge repertoire of antibodies, having infinite types of antigen specificities, from a limited pool of gene fragments encoding the various V(D)J domains. This process is known as V(D)J recombination, and is the hallmark of adaptive immunity [63]. The process of gene rearrangement begins in the heavy chain loci of the earliest progenitor cells that get fully committed to B cell lineage. Initially, μ gene rearrangement starts. If DH to JH recombination is successful, VH to DJH recombination follows. Formation of a productive μ gene on any one of the alleles results in expression of a functional immunoglobulin heavy chain μ (Igμ protein) on the cell surface and differentiation of the cell into precursor B cell (pre-B cell) [64, 65]. Once a successful recombination results in formation of a functional heavy chain, further rearrangements in remaining heavy chain loci cease. This newly formed functional heavy chain is stabilised by pairing with an invariant light chain. This heterodimer forms the pre-B-cell receptor (pre-BCR) in association with the signal transducing units, Igα and Igβ [66, 67]. In the next step, rearrangement in the immunoglobulin κ (Igκ) light chain gene locus begins for generating a successful recombination of its V and J fragments [68, 69]. This step of κ light chain gene rearrangement is stimulated by the newly formed Igμ heavy chain [70]. In case this rearrangement does not succeed, rearrangement begins in the immunoglobulin λ (Igλ) light chain gene locus for recombination of its V and J fragments to give rise to a functional λ light chain [71, 72]. If light chain gene rearrangement is successful, the light and heavy chains combine to form the functional IgM antibody molecule. This antibody is membrane bound and is expressed on surface of the B cell. Only those B cells which express a functional BCR as well as a functional IgM on their surface, progress to give rise to mature B cells.
\nThe process of V(D)J recombination is never perfect. This imperfection allows development of diversity in the antibody structure. However, this imperfection also leads to formation of non-productive recombinations, which can be as high as two-thirds of the total number of recombinations. Several studies have shown that these non-productive rearrangements can be rescued by VH replacement [73, 74, 75, 76, 77]. Each of these gene segments is flanked by conserved sequences, known as Recombination Signal Sequences (RSS). The RSSs consist of a heptamer (having the sequence CACTGTG) and a nonamer (having the sequence (GGTTTTTGT). They are separated by a spacer, which has a length of either 12- or 23-bp [78]. V(D)J recombination follows 12/23 rule. This rule refers to the fact that recombination is preferred between those gene segments that are flanked by RSSs of different spacer lengths. These RSSs are substrates for enzymatic activity of the products of recombination-activating genes (RAG-1 and RAG-2) [79, 80, 81, 82]. This entire enzyme-substrate system is under tight regulation of systems that operate in tissue-specific and stage-specific manner. This ensures that they work in accordance with the lymphocyte developmental pathway [83].
\nTranscription factors (TFs) play vital role during the entire process of B cell development and maturation from HSCs [84, 85, 86]. The entire process involves multiple changes both at the levels of transcriptional state as well as chromatin structure. They include at least four broad areas: (a) developing a localised chromatin state that will be favourable for gene activation through priming the enhancers in lineage-specific manner [87, 88]; (b) expressing TFs in lineage-specific manner [89]; (c) interaction between the various TF networks resulting in formation of extremely complex and fine-tuned regulatory networks that can activate various gene networks in lineage-specific manner [90]; and (d) activating TFs repression mechanisms to prevent development of alternative cell fates so that lineage-specific decisions and commitments do not get altered [91, 92].
\nDuring embryogenesis, three SOXF factors (SOX7, SOX17 and SOX18) play regulatory role in development of haematopoietic system [93]. Out of them, SOX17 is directly involved in the process of expansion of foetal HSCs [94]. A very recent study has shown that SOX7 promotes formation and proliferation of early blood progenitors; and blocks lineage commitment and formation of B lymphocytes [95].
\nThe adult haematopoietic system is established by the coordinated functioning of three TFs, c-myb, acute myeloid leukaemia (AML)-1 and SCL-Tal [96]. Another TF MEF2C is present in very high levels in CLPs and B lymphocytes in the bone marrow [97]. Recently it has been reported that this TF protects B cell progenitors and helps in their survival by enhancing expression of the factors that are involved in DNA repair and recombination [98]. Arid3a and Arid3b, members of the ARID (AT-rich interaction domain) family of TFs, are required for B cell development. However, HSC development can take place independent of Arid3b [99]. Three main TFs, E2A [100, 101, 102], Ikaros [103, 104] and PU.1 [105, 106] specify B lineage commitments in the progenitor populations of HSCs and MPPs. As a result, LMPPs are generated from them. The effects of some of these TFs work in dose-dependent manner. For example, levels of the TF PU.1 in the MPPs determine whether they will progress towards myeloid or B lymphoid lineage [107, 108, 109]. EBF (early B-cell factor) 1 is a known TF that plays vital role in B cell differentiation. In CLPs, E2A has been shown to regulate expression of this TF [110, 111]. In turn, EBF-1 acts in coordination with the TFs E2A and Foxo1 to regulate expression of Pax5 gene, which plays extremely important role in B cell development [112].
\nThus it seems that a regulatory network of several TFs determines and regulates lineage commitment in a recurrent manner [113]. Other significant contributors to this regulatory network include Gfi1 [114, 115], members of NF-κB family [116, 117, 118], members of interferon regulatory factors (IRF-4/Pip and IRF-8/ICSBP) [119], T-bet [120], E47 [121], Krüppel-like factor 3 (Klf3) [122] and Fli-1 [123].
\nEpigenetic control plays a vital role in the entire process of lineage commitment and downstream B cell development. Tet (10–11 translocation family) enzymes, known to oxidise 5-methylcytosine (5mC) to 5-hydroxymethylcytosine (5hmC), are important regulators of somatic cell differentiation. Recent studies have indicated that at least two Tet enzymes, Tet2 and Tet3 are involved in the process of tissue-specific methylation of DNA essential for B cell differentiation [124]. Other studies have indicated the role of high mobility group (HMG) proteins, in particular HMGN, in the process of activation of naive B cells. This group of proteins was reported to act on the chromatin regulatory sites of the resting B cells [125]. Interestingly, the role of various TFs in regulating chromatin accessibility is also being observed. For example, a very recent study on EBF-1 has revealed that its C-terminal domain (CTD) is essential for gaining access to those regions of the genome that are least co-occupied by other TFs. This allows those regions of untouched chromatin to become accessible for structural modulations, such as demethylation. Subsequently, changes leading to B cell fate take place [126]. Earlier studies had shown that Ebf-1 deficient cells get stalled at the point where B cell lineage commitment gets implemented [127, 128]. Gain- and loss-of-function studies on EBF-1 have also shown that this TF upregulates genes that promote B lineage commitment and downregulates those genes that lead to commitment towards non-B lineages [129, 130], thus further confirming the regulatory role played by this TF.
\nB lymphocytes are key players in the immune regulation system. Thus, any alteration in their development or functioning is manifested in the form of a diseased state. Complete understanding of the pathways and the underlying molecular mechanisms of B cell development/functioning is vital as it may lead towards generating new medical interventions. Although we are yet to obtain a clear understanding of the intricacies that govern development of a specific a B cell type, a huge number of studies have contributed towards getting a better picture. Further research is needed for gaining better insight into these processes.
\nNone declared.
Nil.
\nMy sincere apologies go to all those researchers who are doing pioneering work in this field too, but whom I have not been able to quote in my article due to space constraint.
\n"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges".
\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.
",metaTitle:"About Open Access",metaDescription:"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges.\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.",metaKeywords:null,canonicalURL:"about-open-access",contentRaw:'[{"type":"htmlEditorComponent","content":"The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\\n\\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
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\\n"}]'},components:[{type:"htmlEditorComponent",content:'The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\n\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
\n\nAt IntechOpen today, we are still as committed to working with organizations and people who care about scientific discovery, to putting the academic needs of the scientific community first, and to providing an Open Access environment where scientists can maximize their contribution to scientific advancement. By opening up access to the world’s scientific research articles and book chapters, we aim to facilitate greater opportunity for collaboration, scientific discovery and progress. We subscribe wholeheartedly to the Open Access definition:
\n\n“By “open access” to [peer-reviewed research literature], we mean its free availability on the public internet, permitting any users to read, download, copy, distribute, print, search, or link to the full texts of these articles, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, without financial, legal, or technical barriers other than those inseparable from gaining access to the internet itself. The only constraint on reproduction and distribution, and the only role for copyright in this domain, should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited” (reference: http://www.budapestopenaccessinitiative.org)
\n\nOAI-PMH
\n\nAs a firm believer in the wider dissemination of knowledge, IntechOpen supports the Open Access Initiative Protocol for Metadata Harvesting (OAI-PMH Version 2.0). Read more
\n\nLicense
\n\nBook chapters published in edited volumes are distributed under the Creative Commons Attribution 3.0 Unported License (CC BY 3.0). IntechOpen upholds a very flexible Copyright Policy. There is no copyright transfer to the publisher and Authors retain exclusive copyright to their work. All Monographs/Compacts are distributed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0). Read more
\n\nPeer Review Policies
\n\nAll scientific works are Peer Reviewed prior to publishing. Read more
\n\nOA Publishing Fees
\n\nThe Open Access publishing model employed by IntechOpen eliminates subscription charges and pay-per-view fees, enabling readers to access research at no cost. In order to sustain operations and keep our publications freely accessible we levy an Open Access Publishing Fee for manuscripts, which helps us cover the costs of editorial work and the production of books. Read more
\n\nDigital Archiving Policy
\n\nIntechOpen is committed to ensuring the long-term preservation and the availability of all scholarly research we publish. We employ a variety of means to enable us to deliver on our commitments to the scientific community. Apart from preservation by the Croatian National Library (for publications prior to April 18, 2018) and the British Library (for publications after April 18, 2018), our entire catalogue is preserved in the CLOCKSS archive.
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