text:
Physical, biological properties and functions of immunoglobulins.
\r\n\tIn this book Advanced application of radionuclides are introduced. New global trends on safe application of radionuclides in human life is elucidated.
Autoimmune diseases have been increased for the past decades worldwide [1, 2].The prevalence of autoantibodies induced autoimmune diseases is over 2.5% [3].\n
Failure of immunologic tolerance may cause the development of autoimmune response and then autoimmune disease [4, 5].\n
The cause of autoimmune diseases is an association of genetic tendency and environmental factors cause alteration the immune regulatory genes by diver's mechanisms as epigenetics. In autoimmune diseases pathogenesis, both cellular (as in multiple sclerosis) and humoral (as in systemic lupus erythematosus (SLE)) type of the adaptive immune system takes a role. An autoimmune response does not inevitably signify the autoimmune disease [5, 6].\n
In most of autoimmune diseases, the autoantibodies could been found but not all. Even in some autoimmune diseases, the autoantibodies signify not autoimmune disease risk, but also the level of the autoantibodies signifies the severity. By autoantibodies, we can understand immunologic tolerance failure and pathogenesis mechanisms [7, 8, 9, 10, 11, 12, 13, 14, 15].\n
Autoantibodies are self-reactive antibodies. The self-antigens may be found in all cell types (e.g. chromatin, centromeres) and those autoimmune diseases is systemic or be highly specific for a specific cell type in one organ of the body (e.g. thyroglobulin in cells of the thyroid gland) and those autoimmune diseases is organ-specific. The self-antigens can be in proteins, nucleic acids, carbohydrates, lipids structure [16]. Immune tolerance is succeed by various mechanisms, occurred at both central and peripheral organs.\n
An antibody molecule and also autoantibody are include of four polypeptide chains; composed of a pair of identical heavy (H) and light (L) chains. Molecular weight of light chain is 25 kDa and heavy chain is 50–70 kDa. The four chains joint together as a Y shaped. Each light chain is bound to one heavy chain, and the two heavy chains are bound to each other by disulfide bonds between two cysteine amino acid.\n
The antigen-binding site of chains that diverse at different antibody is called as the variable (V) regions and composed of amino acid N-terminal domains of the heavy and light chains. The part next to the V region is called the constant (C) region. A light chain is made up of one V and one C region, and a heavy chain has one V and three (at IgG, IgA) or four (at IgM, IgE) C regions. Each of them is 110 amino acids in length and fields into a characteristic three-dimensional shape called immunoglobulin (Ig). There are three hypervariable regions or CDRs at each variable region of the heavy chain (VH) and of the light chain (VL) which is just 6–10 amino acids in length. CDR3 is the greatest variability of three hypervariable region, at the junction of the V and C regions [4, 5] Figure 1.\n\n
Fab fragment (fragment antigen binding) is composed of a bonded whole light chain (with one V and one C region) and a heavy chain's V and first C region and recognizes the antigen. The Fc fragment (fragment crystalline) is the remaining region of heavy chain. Each antibody contains two identical Fab fragments and one Fc fragment. The hinge region is located in the middle of the Fab and Fc regions and is very bending so helps the two Fab fragment getting closer to antigen far away. The C-Terminal end of the heavy chain of bound antibodies can terminate with or without anchoring in the cell membrane, but the C-Terminal end of the light chain terminates freely without attaching the cell membrane [4, 5].\n
There are two types of light chains according to C region, called κ and λ. Their functions are same. 60% of antibodies are κ chains and 40% are λ chains There are five types of heavy chains also according to C region, called μ, δ, γ, ε and α. Every combination of heavy chain and light chain is available. Antibodies are classified and entitled according to their heavy chains types (IgM, IgD, IgG, IgE and IgA) [4, 5].\n
There is five antibody isotypes with different functions and physical and biological properties, summarized in Table 1.\n
|Heavy chain type\n
|% of total immunglobulin in serum\n
|Monomer or pentamer\n
|Monomer or dimer\n
|Molecular weight (×1000)\n
|170 or 400\n
|Complement fixation\n
|Cross the placenta\n
|Allergic response\n
|Antigen receptor at B cell\n
|Secretoral response\n
1. IgM: Heavy chain type is μ. It has pentamere structure with five Fc fragments where complement binds. The antigen+ pentamere antibody+ complement bound to five Fc complex starts strong complement activation and is removed by phagocytic cells or complement mediated lysis. So IgM plays critical role in neutralization but it has relatively low affinity and cannot penetrate into cells/tissues because of the pentamere structure. Half-life of IgG is approximately 10 days [4, 5].\n
2. IgG: Heavy chain type is γ. It has monomere structure and penetration rate is high e.g. penetare through the placenta There are four classes of IgG: G1, G2, G3 and G4. 65% of total IgG is G1. G1 and G3 activate complement system if the antigen is protein structure and the protein antigens are removed by phagocytic cells. G2 and G4 play role if the antigen is carbohydrate structure. Half-life of IgG is approximately 21 days. Since IgG has high affinity and high molar concentrations in plasma, it makes neutralization [4, 5, 17]. It also makes opsonization because of γ receptors of phagocytes. If N-terminal end is N-acetyl glucosamine, the IgG act as pro-inflammatory and if there is sialic acid then act as anti-inflammatory.\n
3. IgA: Heavy chain type is α. It has monomere or mostly dimere structure which consists of two basic units joined by a J chain. There are two classes of IgA: A1 and A2. IgA1 is in the serum while IgA2 is in secretions as; colostrum, salivary, eye tear, respiratory, digestion and genital and make neutralization of antigens at the mucosal sites. A2; secretory IgA (sIgA) is protected from lytic enzymes in the digestion tract by secretory component (SC) which is a part of the receptor and remains attached to the IgA-dimer [4, 5, 17].\n
4. IgD: Heavy chain type is δ. It has monomere structure. There are two classes of IgD: soluble and bound IgD. While function in immunology of soluble IgD is not known yet, IgD that bound on the cell membrane of newly produced B lymphocytes with IgM, activates of newly produced B lymphocytes by antigens [4, 5, 17].\n
5. IgE: Heavy chain type is ε. It has monomere structure. IgE plays role in parasitic infections and allergic reactions by binding to specific IgE receptors on mast cells and basophiles [4, 5].\n
Antibodies are responsible for the humoral type of adaptive immune responses, glycoprotein structure and produced by B lymphocytes.\n
Antigens can directly bind to antigen receptors of specific B lymphocytes. The type of reversible bond is non-covalent as; electrostatic attraction, hydrogen bonds, Van der Waals-, charge interactions and hydrophobic forces. Membrane–bound antibodies (IgM and IgD type) work as antigen receptors of B lymphocytes (BLR) and can bind to antigens in proteins, lipids, carbohydrates and nucleic acids structures. T lymphocytes can react antigens just in protein structure. For an antigen-presenting cells (APC), there is not any necessity to present antigens to B lymphocytes. Epitopes antigens recognized by T cells are narrow linear peptides from 8 to 20 amino acids [4, 16, 17].\n
After binding of antigens to the receptors that are membrane- bound antibodies; IgM and IgD type, B lymphocyte become activated. The clonal expansion which means proliferation of antigen specific cells follows the activation of B lymphocytes and they differentiate into antibody-secreting effector cells. The specificity of the naïve B cell membrane-bound antibody receptors is same with the secreted free antibodies. During their differentiation period, some B cells may differentiate to produce antibodies with different heavy chain classes (or isotypes) called as heavy chain class (isotype) switching. After switching, different effector functions can be monitored. Repeated exposure to an antigen leads to the production of antibodies with increasing capacity to bind the antigen; called as affinity maturation [4, 18].\n
Antibodies responses are classified into two based on the requirement for T cell help; as T-independent or T-dependent\n
If the structure of antigens is non-protein as polysaccharides, lipids, nucleic acids and others antibody responses evoke without the helper T cells participation. These non-protein antigens cannot bind to MHC molecules consequently cannot be detected by T cells.\n
For immunoglobulin receptor mediated signal transduction in B lymphocytes, the bringing together of two or more antigen molecules in an aggregate (cross-linking), or repeating epitopes of one antigen molecule is needed for antigen binding to membrane bound antibody of the B cell. Multivalent epitope (multiple identical epitopes) as in polysaccharide and lipid antigen can make cross-link many antigen receptors on a specific B cell consequently stimulate proliferation, differentiation and antibody production of B lymphocytes [4, 19].\n
Most soluble protein antigens cannot make cross-link because they do not contain multivalent epitope so cannot stimulate their proliferation and differentiation of B lymphocytes. Antigen-presenting cells process and helper T lymphocytes remember the protein [5].\n
Stimulations of two or more protein antigens lead at least three changes in B lymphocytes to improve the interaction of these B cells with helper T lymphocytes.\n
The changes are:
Increased expression of B7 co-stimulator,
Increased expression of cytokine receptor
Reduced expression of chemokine receptors.
The T cell activation by B cell requires antigen recognition and co-stimulation:
Antigen recognition: B lymphocytes work as antigen-presenting cells (APCs); B lymphocytes may bind, internalize and process the antigen protein, and present multiple different peptides of that protein to T lymphocyte.
Co-stimulation: The helper T cells are stimulated by B7 molecules as co-stimulator expressed by B cells.
CD40 ligand (CD40) and cytokines are expressed by CD4+ helper T lymphocytes after activation. CD40 ligand; a surface protein delivers the co-stimulatory signal in B cells and interacts with CD40 on the surface of B lymphocytes. Attachment of CD40 and cytokines stimulate B cell clonal expression and antibody production. Class switching and affinity maturation are also stimulated by helper T lymphocytes [4, 5, 19].\n
After B lymphocytes proliferation and differentiation into antibody-secreting plasma cells, the antibodies enter the blood through lymphoid follicle. Some plasma cells move to bone marrow, live at the bone marrow for months or years and continue to produce antibodies afterwards antigen is removed. These antibodies supply a rapid response when they meet with same antigen. The humoral immune response decreases physiologically by time because of programmed B cell death. But a small number of activated cells differentiate into memory cells, which "freeze" in a state for a very long time [4, 18, 19]. When the body encounter with the same antigen, the memory cells quickly change into antibody-secreting plasma cells and produce immunoglobulins. The two advantages of memory cells;\n
1. Shorter reaction day: instead of five or more days, it takes one or two.\n
2. B memory cells differentiate with class switch and somatic hypermutation so in case of reinfection, only memory cells with higher affinities and class switch are selected which are completely same with the B cell receptors of the original infection. Recurring antigen stimulation causes to helper T lymphocytes increase consecutively antibody increase with heavy chain class switching and affinity increase [17, 19].\n
Failure of immunologic tolerance can cause the development of autoimmunity. With a genetic background, intolerance can be triggered by environmental factors as sunlight, drugs, chemicals, and infectious agents [5].
Genetic factors: Immunologic tolerance failure is multifactorial and genetic factors are just one of the cause. For example, the relative risk of having autoimmune disease is 5–50 times higher in siblings of affected individuals than in unrelated ones. Multiple genes; mostly MHC predispose to autoimmune disease and genetic predisposition is detected in many autoimmune diseases. For example, individual with HLA-DR4 gene can be suffered from rheumatoid arthritis but not everyone [5, 6].
Environmental factors: İnfections can cause the autoimmune diseases by activating self-reactive lymphocytes. The mechanism is like that an infection lead to a local immune response and activation of APCs. Activated APCs secretes co-stimulators - cytokines and stimulate self-reactive T cells which react with self-antigens in the tissue [4]. Some peptide antigens of microbes are similar to self-antigens, so leads to cross-reactions; called as molecular mimicry [6]. For example; the antibodies against Porphyromonas gingivalis; a periodontal pathogen were increased before RA onset and had a relation with RA [20, 21, 22, 23].
Microorganisms related autoimmune diseases are listed in Table 2.\n
|Related autoimmune diseases\n
|Streptococcus pyogenes\n
|Rheumatoid fever\n
|Escherichia coli\n
|Primary biliary cirrhosis\n
|Shigella spp.\n
|Reiter syndrome\n
|Hepatitis B\n
|Multiple sclerosis\n
|Coxsackie B4\n
|Type 1 diabetes mellitus\n
Sun lights can trigger lupus diseases. Many drugs as procainamide, hydrocarbon pristine, hydralazine, chlorpromazine, methyldopa, quinidine, minocycline and nitrofurantoin can trigger autoimmunity or autoimmune disease through ANAs and ANCAs. Many chemical agents include heavy metals as mercury, gold, and cadmium, pesticides, herbicides, hydrazine can trigger autoimmunity [5].\n
In organ-specific autoimmune diseases, such as thyroiditis, type 1 diabetes mellitus and primary biliary cirrhosis, autoantibodies can be stimulated by infection of the target organ, through molecular mimicry [16, 24]. In systemic autoimmune diseases, such as systemic lupus, autoantibodies can be triggered by genetic factors. For example; a nuclear autoantibodies produced by antigenic drive from excessive release of death cells antigens and enhanced by intrinsic abnormalities in B or T cells [16, 24].\n
The immune system can differentiate self from non-self [5]. Immunologic tolerance is lack of response to self-antigens that encounter with lymphocytes [6]. The recognition of self is a special set of immune events that all constituents of the organism take a role and may be interrupted by environmental and genetic factors [25, 26, 27, 28, 29]. There are three possible immune responses according to antigen type, after antigen encounter with the lymphocytes which has the receptors for a specific antigen;
Active immune response: Due to the active lymphocytes and antigen type is called immunogenic. For example, most of non-self-antigens
Tolerance: Due to inactive or killed lymphocytes and antigen type is called tolerogenic. For example, self-antigens
Ignorance: The antigen cannot either stimulate immunity or induce tolerance. This situation is called as ignorance. For example, self-antigens [4, 5].
Immune tolerance is set of immune events, operating both at central immune organs and peripheral ones.
The immature T cells die by apoptosis, whenever encounter with self-high avidity protein antigens in the thymus. The immature lymphocytes in the thymus can recognize both self and non-self-antigens. If a self-antigen high in concentration and avidity meet with immature lymphocyte, lymphocytes receives signals that trigger apoptosis, finally dies. This is known as negative selection. Since the self-protein antigens are expressed mainly in thymus because of transcription factor responsible called AIRE (for autoimmune regulator), they are high in concentration [4, 19].\n
Some lymphocytes which escape from negative selection, mature to dangerous self-reactive T cells with CD4+ T and CD8+ T. They recognize self-antigens through class I and II MHC molecules [4, 18, 19].\n
And some other develop into regulatory T cells which regulate mostly suppress both naïve and memory T cell responses by a cell to cell contact and by down-regulating the expression of cytokines and co-stimulatory molecules on the antigen-presenting cells. Unfortunately this is not antigen specific reaction [4, 5, 18, 19].\n
Anergy: Anergy is the functional inactivation of T lymphocytes occurs whenever level of the co-stimulators (second signals) is not enough for T cell activation. If level is enough, the co-stimulatory signal which is taken by CD80 and CD86; interaction of molecules expressed on the surface of APC or B cells, reacts with CD28 (or other receptors) on the T cell surface. If T cells with receptors for the self-antigens encounter with sufficient level of self-antigens (signal 1) but do not receive sufficient signal 2, they may induce long-lived T cell anergy [4, 5, 18, 19].
Deletion: Activation-induced cell death: Repeated activation of mature T lymphocytes by repeated encountering with the same antigen cause apoptosis and this is called deletion or activation-induced cell death [4, 18].
Immune suppression: Autoreactive mature T lymphocytes that encounter with self-antigen may develop into regulatory cells which suppress the self-reactive lymphocytes response [4, 18, 19].
If the self-antigens are in structure of polysaccharides, lipids and nucleic acids antigens, they must induce tolerance in B cell and prevent autoantibody production [4]. The B cell tolerance is a set of actions and finally ends with the depletion of or inactive autoreactive B cells. These processes occur at the every stage of B cell [30, 31].\n
When immature B cells encounter with self-antigens in the bone marrow, the B cells are killed and the process is called as negative selection [4].\n
When immature B cells recognize self-antigens in the bone marrow, they may activate their genes of antibodies and start to express a new light chain. These light chains bind to the previously produced Ig heavy chain to produce a new antigen receptor. This process is called receptor editing. The mechanisms of B cell tolerance are multifaceted and may involve receptor editing, controlled migration, and limited availability of BAFF, CD22, Siglec-G, miRNA, and follicular regulatory T cells [30, 31, 32, 33].\n
When mature B lymphocytes encounter with high concentration of self-antigens and B cells producing antibodies that bind with high affinity to self-antigens in peripheral lymphoid tissues, they become anergic; functionally inactivation. T cell-independent antigens can trigger strong signals in the B cell. If it is not strong, the B lymphocytes become anergic [30, 34, 35].\n
Roles of self-reactive B cells are changing according to binding affinities to self-antigens. If self-reactive B cells produce antibodies with high affinity, they undergo elimination or anergy. But if self-reactive B cells produce antibodies with medium or low affinity, they may escape from anergy, even in non-autoimmune individuals [30, 34, 35].Therefore, a significant proportion of immunoglobulins in healthy individuals are made by these autoantibodies. Most of the medium/ low affinity antibodies are multireactive and recognize both self and non-self-antigens [30, 35]. They are called as natural antibodies or natural autoantibodies [16, 17, 36]. Because of their multireactivity, the natural antibodies take an important role in the first part of defense against infections [16, 37] and natural autoantibodies in the development of the B cell repertoire [38].\n
Most of natural autoantibodies are IgM isotype, polyreactive with moderate and low affinity. Therefore, they bind to several unrelated antigens. Also there are natural mono-reactive antibodies [16, 36, 39, 40]. Natural autoantibodies are expressed mostly by CD5+ B1 cell which is the most common B lymphocytes in the neonatal period and in marginal zone B cells [41, 42]. These B1 lymphocytes actively present antigens [43] and also play an important role in the pathogenic autoantibodies production of some autoimmune diseases, as rheumatoid arthritis, Sjögren syndrome, primary antiphospholipid syndrome and systemic lupus [44].\n
In the infantile periods as an evolutionary process, proteins participate mainly in the building and protection of the organism from non-self and self-antigens. During evolution period, these proteins are highly preserved as the autopolyreactive IgM natural autoantibodies (Nabs) produced mainly by B-1 CD5þ cells [25, 41] and also after class switch, polymeric and monomeric IgG isotype antibodies are produced by mostly B2 cells [25, 45].\n
Natural antibodies take critical roles; such as:
Differentiation self from foreign
Recognition of self
At evolution period, autopolyreactivity
First line defense against non-self-antigens; bacterial and viral infections [46].
Regulate the immune system protect the system against tolerance breakdown and the autoimmune diseases.
Maintain tissue balance [47]: Up or down regulation of immunotolerance leads to susceptibility/ progressive or protective role in disease as chronic inflammatory disease [48], cancer [49], cardiovascular disease [50], and certain neurodegenerative conditions [33, 34].
Clearance of tissue and cell debris after degradation [51]; Most diseases is resulting the destruction of tissues/cells which leads to the continuous antigens release. Natural autoantibody recognizes antigens in cell debris and can react with specific antigens of target tissues. In case of chronic inflammation, more natural autoantibody can be stimulated and some autoantigens can mutate to xenoantigens; after these mutations, more specific pathogenic or protective antibodies can be produced [52].
During cell death, some multiple intracellular enzymes as nucleases and proteases are activated which cause the numerous cellular molecules cleavage; as a consequence, some hidden antigens are exposed and called as 'neoepitopes' or neodeterminants. Most of the neoepitopes are undergo to tolerance, but some undergo modification; as cleavage, phosphorylation and oxidation. The self-antigens released by dying cells can be changed by ultraviolet light, oxidation or cleavage by granzyme B [53] delivered by cytotoxic T cells and this change can lead to autoimmune responses. In rheumatoid arthritis, cyclic citrullinated peptides autoantibodies (anti-CCP antibodies) are one of a neoepitope secondary to inflammation [54]. Citrulline is formed by deamination of the arginine amino acid during inflammation/oxidative stress or apoptosis.\n
In specific autoimmune diseases, some of autoantibodies could be detected before beginning of the disease. For example; in SLE, rheumatoid arthritis, type I diabetes, limbic encephalitis and primary biliary cholangitis [55].\n
Changing from preclinical to clinical autoantibody has certain steps. In genetic predisposed individuals, autoantibodies are produced by autoreactive cells. These preclinical autoantibodies can stay for months or even years in these individuals. Under proper environmental conditions, the autoreactive cells would be activated and proliferated. Then, they produce large amounts of autoantibodies and inflammatory cytokines, which lead to tissue injury and the clinical symptoms are observed [6].\n
Natural autoantibodies can provide the templates for the higher-affinity and class-switched pathogenic autoantibodies, under appropriate conditions [16].\n
Production of pathogenic autoantibody:\n
1. Somatic hypermutation: Each antibody can bind at least 2 (IgG, IgD and IgE isotypes) – maximum 10 epitopes (IgM isotype) of an antigen, which has identical epitopes and are close enough. If the multiple antigen-antibody bind each other, the total strength of the bond is much greater than a single one. This is called the avidity of the interaction. The molar concentration of an antigen needed to occupy half the available antibody molecules in a solution is the dissociation constant (Kd) and used for expression of affinity. The lower the Kd means the higher the affinity. In a primary immune response, produced antibodies have a Kd in the range of 10−6–10−9 M and after encountering with repeated antigens, the affinity can rises up to 10–11 M. This increase in antigen-binding strength is called affinity maturation or somatic hypermutation [4]. Mostly point mutations in the genes responsible for variable regions of antibody are detected [16]. They happen in the germinal centers of secondary follicles and AID enzyme that initiate them [17].\n
2. Class switching: The membrane bound IgM and IgD the antigen receptors of naïve B lymphocytes. After stimulation, the antigen specific clone B lymphocytes may proliferate and differentiate into antibody-secreting cells. Some of these B cells may secrete IgM, and some others may produce antibodies of other heavy chain classes. The change in Ig isotype production is called heavy chain class switching. The V regions remains same, specificity of B cells maintains [4].\n
The exons encoding the constant regions of all antibody classes on chromosome 14, are placed with μ (for IgM) nearest to variable region segments, followed by γ (IgG), α (IgA) and ε (IgE). By a successful VDJ rearrangement, first the nearest constant region which is μ is used, resulting in the production of IgM [17]. Unmutated or minimally mutated recombined VDJ gene sequences encode the multi and monoreactive natural IgM antibodies/autoantibodies [56]. AID deaminates cytidines in immunoglobulin VDJ and switch-region DNA, then ssDNA nicks, gaps or double-strand breaks are generated. Repair of these lesions involving error-prone translesion DNA polymerases are made by the B cell DNA and this results in insertions of point mutations or resolution of double-strand breaks, and hence, class-switch DNA recombination [57]. After class switch with the same variable region, these cells can express IgG if the exons encoding the γ constant region; IgA if it is α constant region; and IgE if it is ε constant region. T-lymphocytes and other cells release cytokines influence isotype of class switch [17].\n
Unmutated natural IgM autoantibodies expressed by B1 cells provide the 'templates' for the high-affinity and class-switched IgG and/or IgA autoantibodies which can cause autoimmune diseases [49, 58, 59]. Anti-DNA, anti-insulin and anti-IgG (RF) autoantibodies are pathogenic high-affinity autoantibodies that undergo somatic hypermutation, class-switch DNA recombination and antigen driven clonal selection detected at systemic lupus, type 1 diabetes and rheumatoid arthritis patients [60]. Somatic hypermutation and class-switching [56, 60, 61] including the expression of activation-induced cytidine deaminase (AID) [62] are associated with the expansion of B-2 cells.\n
Class switch and somatic hypermutation are initiated by the same enzyme, AID, in the germinal centers of secondary follicles parallelly [17].\n
3. Somatic diversity: Somatic recombination: Antibodies are capable of binding a wide variety of antigen, since variable region of antibody molecules forms a flat surface field into different shapes. The epitopes or determinants are the parts of antigens that are recognized by antibodies based on sequence (linear determinants) or shape (conformational determinants). Some hidden antigen molecules are exposed after a physicochemical change, called as neodeterminants [4].\n
Diversity of antibodies is generated by the genetics arrangement of antibody production; unique molecular random generator. The variable region of an immunoglobulin is formed by both the heavy and the light chain which are carried on different chromosomes [5]. The variable portion of the heavy chain is encoded in separated gene segments of three types, V (variable; the number of gene segments is 65), D (diversity; 27) and J (joining; 6). A complete heavy chain variable region exon is randomly cobbled together by juxtaposing one V, one D and one J segment by a cut and paste process at the DNA level by an enzyme complex containing RAG-proteins (recombination activating gene) which excises intervening DNA, and normal DNA repair proteins directly rejoin the segments. Light chain genes have just V and J segments, not D [17]. In summary, the diversity of antigen binding is achieved by mostly V genes and their combination with different D and J genes. Different antibodies are produced by four different mechanisms as; randomly combining V-(D)-J segments, randomly combining heavy and light chain, imprecise joining and somatic hypermutation [4, 17]. Somatic diversity is performed during central B cell intolerance.\n
4. Genetic abnormalities: Some genetic alterations results clinical autoimmune disease but some alterations are influenced by environmental factors. For example; single gen knockout and overexpression lead to clinical autoimmune disease while most of the autoimmune disorders are polygenic. Three examples of spontaneous or induced genetic alterations lead to clinical diseases [16].
Abnormal survival of autoreactive lymphocytes: Mutations in Fas/CD95 causes over expression of the B cell stimulator BLyS; BAFF and the antiapoptotic regulator Bcl-2 which leads the abnormal survival of autoreactive lymphocytes. It causes an autoimmune lymphoproliferative syndrome/Canale Smith syndrome in humans [16, 63],
Defective removal of apoptotic cells: A group of proteins as Mer and serum opsonins (e.g., natural IgM antibodies, C1q, serum amyloid P component [SAP] and milk fat globulin epithelial growth factor-8 [MFGE8]) [64] take role in the removal of apoptotic cells. In Mer deficiency, macrophages take a proinflammatory signal not an anti-inflammatory one for ingestion of apoptotic cells. If there is a defective clearance of apoptotic cells in surface IgM, C1q, SAP and MFGE8, clearance of apoptotic cells leads to postapoptotic necrosis and/or through lack of engagement with specific inhibitory receptors on the phagocyte. In MFG-E8 deficiency, apoptotic cells accumulate in germinal centers and in C1q-deficiency, apoptotic cells accumulate in the kidney. These deficiencies cause lupus-like diseases [16].
Breakdown in the regulation of B cell or T cell activation threshold: If threshold regulators of cbl-b, PD-1 and Zap-70 and the SLAM cluster in T cells, and Lyn and FcγRIIb in B cells change genetically, failure of peripheral immune system could happens. If lymphocytes are more easily activated, they produce more auto-antibodies as in systemic lupus. Mutations of Zap-70 lead to production of RFs as in rheumatoid arthritis [16, 65]. PD-1-deficiency causes lupus in C57BL/6 and myocarditis in BALB/c.
There is some signature autoantibodies cause autoimmune diseases as anti-endomysial antibodies (EMA), anti-gliadin antibodies (AGA). But there is not a specific antibody detected yet in several autoimmune diseases, as psoriasis [6].\n
Lymphocytes and APC are strongly activated by type I interferons (interferon-α and β) [66]. Patients with systemic lupus have elevated levels of interferon and autoantibodies as anti-DNA and Sm/RNP. By binding to chromatin which contains DNA or to Sm/RNP which contain small nuclear RNAs, they enter cells through the FcγR or B cell receptor. The intracellular Toll-like receptor is activated by nucleic acid which leads to production of interferon and activation of immune system. The protein antigen stimulates T cells, probably are responsible for the specificity of the immune response. These are called Toll hypothesis [67].\n
Autoimmune disease can be classified as systemic or organ specific. Systemic autoimmune diseases (Table 3), involve multiple organs or tissues, whereas organ specific autoimmune diseases (Table 4), involve a single organ or tissue. Almost all organs can be affected by either systemic or organ-specific autoimmune disease [5].\n
|Organ(s) involved\n
|Systemic lupus erythematosus\n
|Joints, skin, nervous system, kidneys, blood cells, heart, lungs\n
|Anti dsDNAb\n
Anti Sm b
Anti ribosomal P b
Anti RNA helicase
|Rheumatoid arthritis\n
|Joints, blood, vessels, lungs\n
|Anti citrullinated peptides b\n
Rheumatoid factor
|Sjögren's syndrome\n
|Exocrine glands (salivary and lacrimal glands), kidneys, nerves\n
|Anti Ro60 (SS-A)\n
Anti Ro52
Anti La (SS-B)
|Skin, blood vessels, GI tract, lungs, kidneys\n
|Anti topoisomerase I b\n
Anti fi brillarin (U3 RNP) b
Anti RNA polymerase I b
Anti RNA polymerase III b
|Muscles, lungs\n
|tRNA synthetases\n
(Histidyl, alanyl, threonyl, glycyl, etc.) b
Signal recognition particle b
|Organ(s) involved\n
|Hashimoto's thyroiditis\n
|Thyroid peroxidase\n
|Graves' disease\n
|Thyroid-stimulating hormone receptor\n
|Addison's disease\n
|Adrenal glands\n
|Type I diabetes\n
|Pancreatic islet cells\n
|Glutamic acid dehydrogenase,\n
islet cell antigens
|Pemphigus vulgaris\n
|Desmoglein 3\n
|Bullous pemphigoid\n
|230 kDa hemidesmosomal antigen\n
|Skin melanocytes\n
|Unknown melanocyte antigens\n
|Goodpasture's syndrome\n
|Kidneys, lungs\n
|Type VII collagen\n
|Myasthenia gravis\n
|Nervous system\n
|Acetylcholine receptor\n
|Multiple sclerosis\n
|Nervous system\n
|Unknown myelin antigens\n
|Pernicious anemia\n
|Gastric parietal cells\n
|Parietal cell antigens,\n
intrinsic factor
|Primary biliary cirrhosis\n
|Bile ducts\n
|Dihydrolipoamide acyltransferase and other antigens b\n
|Autoimmune hepatitis\n
|Smooth muscle antigens (F-actin)\n
The antibodies' Fab regions bind to antigens and can block/stimulate the effects of them and the Fc regions can bind to many cells of immune system as phagocytes and complement and activate diverse effector mechanisms to eliminate these antigens; Fcγ-R (for IgG), Fcα-R (for IgA), Fcα/μ-R (for IgA and IgM), Fcε-R (for IgE). The effective binding of antigen-antibody occurs after recognization several IgG molecules. The affinity of the binding is too low with a single, free antibody. Bigger immune complexes by antigen and several Fc parts of antibodies causes to rapid internalization for phagocytosis and antigen clearance. Heavy chain class switching and affinity maturation enhance the protective functions of antibodies. There is an exception to this rule in mast cells and eosinophils, just binding a free (meaning non-antigen-complexed) IgE is enough because of their high-affinity Fc-ε-receptors [4, 17].\n
1. Neutralization of foreign and self-antigens: Antibodies bind to block, or neutralize the activity of foreign or self-antigens [4].\n
2. Opsonization and phagocytosis: Complex of antibodies with foreign and self-antigens promote their ingestion by phagocytes (opsonization). When IgG1 and IgG3 isotype antibodies bind to a foreign or self-antigen, their Fc regions bind to a high affinity receptors called FcγRI (CD64), which are on neutrophils and macrophages. The binding of antibody Fc tails to FcγRI results in opsonization of antigenic molecules into a vesicle called a phagosome, where fuse with lysosomes and activates the neutrophil or phagocytes. The activated ones produces in their lysosomes, large amounts of reactive oxygen intermediates, nitric oxide, and proteolytic enzymes, all of them together destroy the ingested antigenic cells [4].\n
3. Antibody-dependent cellular cytotoxicity (ADCC): Natural killer (NK) cells produce an Fc receptor called FcγRIII, which binds to IgG antibodies. The activated NK cells discharge their granules, which contains proteins that kill the opsonized targets [4]\n
4. Activation of the complement system: Antigens without antibody, as part of innate immune response to infection, and antigens with antibody, as part of adaptive immunity can activate the complement system. The complement system takes role in the elimination of opsonized antigens [4]. Examples; activation of complement causes diseases at kidneys of systemic lupus and lupus nephritis patients, fetal loss associated with the antiphospholipid syndrome [68, 69], autoantibody administration into the transgenic K/BxN mouse of rheumatoid arthritis [70], in glucose-6-phosphate isomerase patient. In the NZB/W F1 murine model of immune-complex-mediated lupus nephritis, mice lacking the FcγRγ chain were protected from nephritis, indicating a critical role for FcγRs in tissue inflammation [71].\n
5. Mucosal immunity.\n
6. Pro-inflammatory and anti-inflammatory effect: natural polyautoreactive IgM antibodies can protect from autoimmune diseases [30]. Also IgG isotype autoantibodies has an anti-inflammatory capacities, according to their IgG subclass and the extent of glycosylation/sialylation of the Fc glycan linked to Asn297 [71, 72]. These properties regulate the binding of antibody to a different Fc-receptors [72]. The receptors as FcγRI (CD64), FcγRIIIA (CD16a), and FcγRIIIB (CD16b) mediate activating signals, but also FcγRIIA and FcγRIIB (CD32) mediate inhibiting signals. Glycosylated/ sialylated different IgG isotypes antibodies bind to Fc-receptors for activating and inhibiting with different affinities [72]. According to glycosylation/sialylation patterns and IgG subclass determine, an autoantibody produces FcγR-mediated either pro- or anti-inflammatory functions [73]. So glycosylation of autoantibody can be an important regulator of autoimmune disorders [74]. While IgG isotypes produced with T cell-dependent reactions were poorly sialylated causes pro- inflammatory, a high degree of sialylation that mediates anti-inflammatory properties [75]. Activated B cells and plasma cells regulate both T cell differentiation into follicular helper T cells and cytokine profiles [76]. By stimulation of TLR, B lymphocytes produce different cytokines to dendritic cells [77]. Dendritic cells are the most important antigen-presenting cells to T cell. B cell also present the antigen to T cell and so promote the proliferation of activated T lymphocytes, the development of robust T effector responses, and normal T cell memory compartments [78]. TLR-signals in murine B cells promote IFN-γ production from T cells and control antibody isotype switching to IgG2 in vivo [77]. The cowork of activated B and T cells is crucial for the antibody responses and their outcome as pathogenic potential, that is, the antibody class and glycosylation/ sialylation pattern.\n
Testing of autoantibodies is diagnostic criteria in many diseases. But, also autoantibodies could be detected in healthy individuals [79]. Since isotype/subclass and glycosylation pattern is critical for the pathogenic potential of a particular antibody, it could be helpful for the diagnostic analysis. Pathogenic autoantibodies could be produced either by continuous formation of short-lived plasma cells or through the formation of long-lived plasma cells, or both [80]. Therapeutic treatment available nowadays could suppress B cell activation and short-lived plasma cell, while do nothing to long-lived plasma cells [81].\n
By contrast, mice with FcγRIIb knocked out spontaneously develop a lupus like disease [71]. Different isotypes antibodies have different affinities for the four FcγRs. IgG2a has higher affinity for FcγRIV, leading to inflammatory responses, whereas IgG1 selectively engages FcγRIIb, leading to inhibitory responses [30]. There is a similar relationships with human FcγRs and that the ability to protect or induce inflammation will change according to the isotype of the autoantibody and FcγR engaged.\n
7. Removal of cell debris: Natural autoantibodies takes role in the removal of cell debris during inflammation, and autoantibodies to inflammatory cytokines have protective functions against inflammation [82].\n
Immune responses can cause tissue injury and disorders called as hypersensitivity diseases. Hypersensitivity is a term of excessive or aberrant immune responses [4]. Tissue damage in autoimmune diseases can occur through several mechanisms, which are similar to three of the classical types of hypersensitivity reactions [5]:\n
1. Type II (caused by autoantibodies reactive with cell surface or matrix antigens):\n
Antibodies against cell and tissue may cause tissue and disease. IgM and IgG antibodies activate the phagocytosis of cells by binding to complement and Fc receptor- mediated leukocyte [4]. The reactions are caused by antibodies against self-protein antigens. Autoantibodies generated against cell surface antigens/extracellular matrix proteins may be cytotoxic (type IIA) or agonistic/antagonistic (type IIB). Autoantibodies to cell surface antigens may initiate cell destruction by complement- mediated lysis (cell destruction), phagocytosis, or antibody-dependent cell-mediated cytotoxicity (ADCC) [5]. At Table 5, some examples of antibody-mediated diseases are given.\n
|Target antigen\n
|Pemphigus vulgaris\n
|Proteins in intercellular junction of epidermal cell\n
|Antibody-mediated activation of proteinase, disruption of intercellular adhesion\n
|Autoimmune hemolytic anemia\n
|Erythrocytes membrane antigen\n
|Opsonization and phagocytosis of erythrocytes\n
|Myasthenia gravis\n
|Acetylcholine receptor\n
|Antibody inhibits acetylcholine binding\n
2. Type III (caused by immune complexes):\n
Autoantibodies can bind to circulating antigens and form immune complexes that deposit in vessels, tissues and cause tissue injury. Injury is mainly due to leukocyte recruitment and inflammation [4]. Autoantibodies can cause disease by forming immune complexes with the circulating antigens. Immune complex formation is a normal process to remove antigens and to phagocyte through Fc or complement receptors so are prevented their deposition. The efficiency of uptake of immune complexes by either Fc receptors or CR1 is proportional to the number of IgG molecules associated in the complex [5]. At Table 6, some examples of immune complex mediated diseases are given.\n
|Target antigen\n
|Systemic lupus erythematosus\n
|DNA, nucleoproteins\n
|Complement and Fc region mediated\n
|Polyarteritis nodosa\n
|Hepatitis B surface antigen\n
|Complement and Fc region mediated\n
|Poststreptococcal glomerulonephritis\n
|Streptococcal cell wall antigen\n
|Complement and Fc region mediated\n
3. Type IV (delayed-type hypersensitivity, mediated by T cells):\n
T cell-mediated disease is caused by CD4 T lymphocytes or by killing of host cells by CD8 CTLs [4]. T cells recognize protein antigen-presenting cells in the context of class II major histocompatibility complex (MHC) molecules and produce the cytokines interferon γ (IFN-γ), interleukin 3 (IL-3), tumor necrosis factor (TNF) α, TNF-β, and granulocyte-macrophage colony-stimulating factor (GM-CSF). Elaboration of "TH1 (a subset of helper T cells) cytokines" leads to macrophage recruitment and activation, enhanced expression of adhesion molecules, and increased production of monocytes by the bone marrow [5]. At Table 7, some examples of T cell-mediated diseases are given.\n
|Target antigen\n
|Rheumatoid arthritis\n
|Antigen in joint synovium\n
|T cell mediated\n
|Type I diabetes mellitus\n
|Islet cell antigen\n
|T cell mediated\n
Forest is a conditional renewable resource which can be regenerated but needs a certain period of time to maintain its sustainable functioning. In India, the forest resources have been found to be depleting at a pace which is much high [1]. Rapid industrialization, urbanization and over-exploitation have resulted not only in decline but also in permanent loss of forest cover to an alarming rate [2]. The major driver behind all these factors is the uncontrolled population growth of humans which leads to the dramatic increase in the demand for wood and forest products. The over-exploitation of forest resources has taken place beyond the sustained yields to fulfil the needs of humans, thus bringing a change in the net forest cover [2, 3]. With the current rate of population explosion, the world population could be expected to increase from 7.6 billion to about 10 billion in the next 30 to 35 years [4]. The growing demand for food can be expected to rise by 50% in the given period, and it is a matter of serious concern. Rational utilization and proper management of the forest resources are the most viable ways to prevent mass destruction of forests and large-scale species extinction. It is necessary to find the links between the growing demands and meeting the demands in a sustainable manner. The scope of future studies must focus on the solution to establish this link by incorporating the principles of forestry, restoration ecology and natural resource economics.
Deforestation occurs when a land initially dominated by naturally occurring tree species is converted to provide certain services in response to the human demand. The UN Food and Agricultural Organization (FAO) has defined deforestation as the conversion of forest to another land use or the long-term reduction of tree canopy cover below the 10% threshold. Forest areas around the world are majorly cleared for agriculture, logging, mining and large-scale developmental projects. The Food and Agricultural Organization [5] coordinated the Global Forest Resources Assessment (FRA) which reported a 3.16% decline in the global forest cover from 1990 to 2015, and the total forest cover stands at about 30.6% in the present time compared to 31.6% in 1990. The rate at which the forest cover is declining poses a direct threat in the near future if not checked. With an estimated annual loss of 18.7 million acres, it is evident that future demands on forest resources would certainly lead to immense competition among nations [6]. Recently, in 2016, a study from the Maryland University reported that 73.4 million acres of the global tree cover were lost. Such a destruction of this essential and self-sustaining resource puts the implementation of the principle of sustainable development as mentioned in the Brundtland Report and Sustainable Development Goals of the 1992 of Rio Earth Summit in the state of question. And, it is an urgency to conserve the forests of which a vital part is already lost.
The value of forest is simple to understand but sometimes tough to quantify. The various values of the forest have been shown in Figure 1. Forest has a major contribution on the global economy and supports livelihood of the majority of rural populations in the world [7]. The direct uses of forest are most easy to quantify as it is directly related to economic returns. The indirect use and option value although play a major role in defining the valuation of the forest resources are seldom quantified and are being intangible in nature [8]. There is also a no-use value of the forest resources which considers leaving the current forest intact as a heritage for the future generation and for satisfaction and happiness of the current population. But, these eco-centric views alone cannot suffice the support for a change in policy and land use pattern. There are some other important values of forests that are difficult to quantify. One of the major roles that forests play is that it acts as a major carbon sink [9, 10, 11]. Plants utilize carbon dioxide in the process of photosynthesis and store it in the form of carbohydrates, and these carbohydrates reach the soil as dead organic matter and contribute to soil carbon sink. When forests are cleared, less CO2 is absorbed by plants, and atmospheric CO2 concentration increases with the passage of time due to unavailable sink. Also, there is a marked reduction in soil organic carbon with the loss of vegetation cover, thereby affecting the productivity of the ecosystem. Productivity is believed to be an indicator of carbon assimilation capacity, and hence the more productive the forests are, the more carbon it stores [12, 13, 14]. The tropical forests are among the most productive as well as the most threatened ecosystem in present time with the maximum rate of deforestation. Thus, it is imperative to control the rate of deforestation in order to avoid the adversities associated with it.
The value and functions of forests are immense and diverse. Similarly, the impacts of deforestation are global and commune the whole forest community. One of the major constraints in understanding the true impact of desertification on forests is the inadequacy and imprecise form of data.
The effect of large-scale deforestation is global, but it is necessary to assess its impacts on microclimate, regional climate and global climate to form a holistic understanding of mechanism [15]. The association of deforestation with the increased CO2 concentration in the atmosphere and changes in the mass balances and surface energy can result in climate change at the local and global level [16]. The change in land use pattern especially the clearing of forest cover affects both hydrometeorological and global CO2 concentrations leading to more warming as CO2 readily absorbs infrared radiation [17]. Clearing tree cover and vegetation leads to increase in albedo of the region as bare soil reflects more solar radiation than vegetation, which again is a factor for altering regional radiation flux [15, 18]. One of the noticeable changes in regional climates occurs when the cloud formation shifts to higher elevations from lowland plains as a result of deforestation in the later area [19]. In a global scale, deforestation leads to warmer and drier weather due to the synergistic effect of reduced evapotranspiration, increased albedo and CO2 concentration that triggers desertification, loss in biodiversity and melting of polar ice caps, ultimately leading to food insecurity. All these effects are successors of extremes in climate variation that are produced by the large-scale reduction in forest cover [20]. The estimated quantity of CO2 added to the atmosphere due to deforestation in the tropics is roughly two billion tonnes [21]. It is interesting to note that the CO2 emissions due to clearing of forest will almost be equivalent to 25% of what is added to the atmosphere due to anthropogenic emissions [22]. The shift in climate is somehow correlated to reducing forest cover. Further researches can clearly define the mechanisms and pathways by which these shifts are progressing and how they can be mitigated.
The global water cycle depends on the amount and distribution of precipitation for which one of the influencing factors is evapotranspiration [23]. There is a direct effect on drinking water on the basis of quality and quantity, fisheries and aquatic habitats, occurrence of flood and drought, life of dams on account of increase siltation and agriculture due to poor quality irrigation and crop yield [24, 25]. It must be recognized that the protective role of forests is operative and has a major impact on urban water resources [26]. Forests play an important role in maintaining the watersheds [27]. The degraded or degrading watersheds can be recovered by forestation, but once the forest or vegetation cover is lost, the watershed becomes vulnerable to erosion. This erosion leads to siltation in the downstream areas and consequently reduces the depth of river bed increasing the chances of flood. There are two main effects of deforestation that increase the chances of flooding. One is by reducing the tree fountain effect, i.e. interception and moisture uptake by the trees would cease after deforestation reducing the moisture holding capacity of soil that leads to greater runoff and erosion. And other is by the process of soil compaction and poor soil structure that will lead to reduced organic matter content of soil devoid of vegetation cover [28]. There are severe long-term effects on soil as a cause of deforestation. During slash-and-burn or shifting cultivation, an area of forests clears and exposes the bare soil to weather extremes of high solar insolation and heavy rainfall [29]. In the absence of the forest cover and organic matter, soil could not accommodate heavy precipitation, and the fertile layers of soil used to be easily washed away ultimately reduces the long-term productivity. The effect on soil is dependent on the interrelation and synergistic effect of evapotranspiration and infiltration that are directly altered due to loss in vegetation cover [23].
Deforestation directly increases erosion and siltation rates. There is an increased risk of flooding in Yangtze River basin in China and the major river basins in East Asia and the Amazonian basin [23, 30]. The slopes and terrains are more vulnerable to such situations. There are formations of shallow gullies which may be accounted for the concentrated flow of runoff that prevails due to long-term erosion. Cultivation and ploughing along the slopes cause rapid erosion in the areas with less vegetation cover [31]. In the Loess Plateau, the slopes of steepness greater than 15° showed shallow gully erosion as a result of cultivation activities [32]. The availability of dead vegetation can reduce the surface runoff from the early season rain and check soil erosion [33]. It is necessary to sustain the forest and vegetation cover to maintain the soil productivity and water quality of both the surface and underground sources.
Forests are very rich in biodiversity and store a vast gene pool, and the majority of species occur in the tropical forests. It consists of two-thirds of all known species and 65% of 10,000 species that are recognized as endangered by the International Union for Conservation of Nature (IUCN) [34]. The biodiversity could be regarded as an important asset that is necessary to conserve for future utilization. The World Health Organization states that the traditional people, almost 80% of them, rely on the local biodiversity for traditional medicines. A loss in the biodiversity may directly affect their health care and well-being [35]. Another most noticeable impact of deforestation is the increasing human-wildlife conflict. The decreasing forest cover is limiting the habitats of many species due to which is forcing them to intrude with human welfare [36]. There are increasing instances of invasion and animal killing. The northern margin of West Bengal, India, forms a significant portion of the Himalaya Biodiversity Hotspot. The area has observed heavy fragmentation in the last decade, and as a result, there was a huge loss in the agricultural crop and both human and elephant life due to conflicts. There are reports of mortality of 20 elephants and 50 persons annually from this area [37]. It is also estimated that if deforestation in the Himalayas continue at the current rate, the dense forest cover (>40% canopy cover) will be restricted to 10% of land area in the Indian Himalayas by 2100. This may lead to a significant loss of 366 endemic plants and 35 endemic vertebrates [38]. The loss in biodiversity is of global concern irrespective of regional and local importance. Conserving the forest and an increase in forest cover certainly find a positive correlation with the increase and sustenance of biodiversity. Conversion of forest land to agricultural fields and other land use could be a threat to major loss in biological diversity in the near future.
Forests contribute to the world economy in terms of timber production and other forest produces. There are different contributions of forest as a means of direct employment in forestry services and other value-added contributions as recreation and aesthetics. The loss of tropical forest cover annually may account for about 45 billion US dollars [39]. The destruction of forest eliminates the sources of economic gain directly obtained and also eliminates the potential gain from the resources that the forest sustains as biodiversity, soil and water. Also, the destruction of forest increases the negative externalities in the form of increasing CO2 concentration, risk of flood and human-wildlife conflict [40]. The proximity of settlements to protected areas further intensifies the chances of human-wildlife conflict (HWC). It is reported that settled households face high risk of HWC due to their close proximity of the Kanha National Park in Central India [41]. Deforestation has its social influence in the form of long-term effect. Development is a serious concern for indigenous community as it certainly leads to a change or shift of their culture and tradition. The cultural and religious aspects of the community are seldom preserved amid infrastructural advancement that leads to land and social conflict [42]. In a study of household survey from rural areas of Madhya Pradesh and Chhattisgarh state of India, it was found that the poorest of the local community gained about 30% of their living from forest produce which was claimed to be even higher than the returns from agriculture. Also, forest provides an option as safety net during the period of crisis in rural areas [43]. Various ecological services provided by the forests have been lost due to deforestation which in turn has immediate effects on the local communities dependent on these services for their daily needs. The inclusive approach for the sustainable management of forest resource is a vital consideration that considers economy as a subset of the society which in itself is a subset of the environment. Such considerations can help sustaining the constantly declining forest cover and its long-term impacts.
The forest cover was found to be increased by ca. 1% for the year 2017 which is 21.54% of the total geographic area when compared to that of forest cover status in 2015 which is a positive sign towards the constant efforts that are being put in to protect the forest (Figure 2). This positive change in the forest cover is mainly attributed to the conservation and management practices that include afforestation activities, participation of local peoples for better protection measures in plantation areas and traditional forest areas, expansion of trees outside forest, etc. Also, with this increase in the forest cover, the country has procured 8th position among the top 10 countries reporting the greatest annual forest area gain. Although there has been an increase in the total forest cover in India, still there are certain regions within the country that has sought a reduction in the same (Figure 3). The main reasons attributed for this reduction are shifting cultivation, rotational felling, other biotic pressures, diversion of forest lands for developmental activities, etc. [44]. A transition in the various forest cover classes has also occurred over the past. The present status of forest cover (%) of India belonging to various forest cover classes has been shown in Figure 4. It has been found that there is a reduction in the moderately dense forest and an increment in the open forest depicting degradation of forest cover to some extent. Changes within the forest cover classes result in decrease in the production capacity, thereby leading to forest degradation [45]. Control and regular check of these activities can help in strengthening the conservatory efforts of forest protection. This is the necessity of the present time to conserve the forests in order to sustain the vital ecosystems and the major carbon sink to combat the effects of global climate change and ultimately maintain the environmental balance.
Forests are an invaluable resource that is being subjected to so many threats. In order to protect our forests, it is very much necessary to understand the reasons behind their destruction. Differentiating the agents and causes of deforestation will enable in understanding the same [36]. Several disturbances within the forest directly or indirectly contribute in destruction of the forest. This can be interpreted from the results obtained while surveying in different forest patches in India.
Arunachal Pradesh is one of the states that has more than 75% forest cover and has the maximum very dense forest cover type [44]. The state is highly diverse in terms of climate as well as forest cover with tropical, sub-tropical, temperate and alpine forests having higher NDVI (≥7.0) [46]. However, the pressure on forest resources is consistently increasing with the rise in population, development activities, large number of wood-based industries and unsustainable land use practices resulting in their degradation [47]. This has also resulted in decrease in the forest cover of the state [44]. Several disturbances were being observed during the field survey in the forests of Arunachal Pradesh during 2007 to 2010. The major disturbances that were found include lopping, cut stumps, litter collection, soil removal, grazing, fire, NTFP collection and fuelwood collection (Figure 5). Of these, fuelwood collection was found to be the most recurrent activity followed by grazing. Generally, fuelwood collection has not been considered as the major cause of deforestation but leads to the same in certain regions with reduced forest area such as in the Philippines, Thailand and parts of Central America [36]. Forest fire has also been observed as an occasional event in certain parts of Arunachal Pradesh. Fires are generally used as a tool in clearing the forest for shifting cultivation which is one of the major agricultural practices performed in the state. Fires when used responsibly act as a valuable tool in managing forest and agriculture, but when abused, it can lead to deforestation [48, 49]. Other disturbances that can be an indicator of deforestation include NTFP collection and presence of cut stumps in certain forests. Forests of the state are highly diverse in endemic as well as nonendemic species, which need intensive monitoring and management to conserve the species-rich ecosystems from ever increasing anthropogenic pressure and changing climatic conditions [50].
Madhya Pradesh is among one of the states of India which is endowed with rich and diverse forests and comprises the largest forest cover in the country [44]. This is mainly because of the efforts that the state has put in to conserve and harness this invaluable resource through innovative measures like community participation and decentralization (MP) [51]. Even after these continuous efforts, there are certain regions within the state where the occurrence of several types of disturbances in the forest has been found. The common disturbances that were being observed during the field survey (2017–2019) include fire, grazing, fuelwood collection, forage removal, litter collection, NTFP collection, lopping, thatch collection, root collection, soil removal, etc. (Figure 6). Among all these fire has been found as the major recurrent type of disturbance in the forests of Madhya Pradesh. Other major disturbances were grazing followed by fuelwood collection. Since every type of deforestation is not intentional but some which are the results of amalgamation of anthropogenic and natural factors like wildfires and subsequent overgrazing can prevent the growth of young trees [52] and thus eventually degrade the quality and productivity of the forest.
The name of the state 'Jharkhand' itself connotes 'area of land covered with forests' and has been exhibiting a unique relation with forests since time immemorial [53]. During the forest cover assessment [44], a net increase of 29 sq. km in the forest cover has been observed in the state which is mainly because of the plantation and conservation efforts within recorded forest areas. Although, there was an increment of 314 sq. km in the forest cover within the recorded forest areas, because of the felling of trees outside the forests area, its effect on forest cover has been offset. Also, several types of disturbances can be seen within the forest areas during the field survey (2016–2018), and the major disturbances were fuelwood collection, grazing, forage removal, lopping, cut stumps, thatch collection, root collection, soil removal, litter collection and NTFPs collection (Figure 7). Most of these disturbances were occasional in nature. Although these disturbances are not that recurrent, a regular check is necessary in order to prevent the forests from degrading and in achieving a sustainable forest cover. The forest management strategies should focus on the increasing demands of different timber and non-timber forest produce to conserve the plant diversity of the natural forests of the state [54].
Forest management and protection by the local communities is an age-old practice in India which can be traced back to the protective nature of the Bishnoi Community of Rajasthan towards the local forest and animals as the black buck. The idea of community-based forest management emerged in an administrative level in the 1970s and 1980s. The declaration of the Government of India in June 1990 marked the establishment of Joint Forest Communities in different India states as per the National Forest Policy of 1988. The Earth Summit of 1992 provided with a clear objective of Sustainable Forest Management to which India responded in a positive way. From an increase in the forest cover, non-timber forest product (NTFP) to conservation of native flora and fauna, a whole new realm of forest management strategy by the collaboration of forest departments and local communities aided in decelerating the degradation of natural forest in India [55]. The Participatory Forest Management (PFM) is equivalent to an informal contract in which the local communities are allowed to consume a portion of harvest and NTFP if they protect and conserve it for 5–10 years. In India there is no legal authority of the local community on the forest resource where as in other countries as in Nepal the Community Forest User Groups (CFUG) are registered under their Forest Act, 1993 [56]. It is the positive effort of the local communities of India that the area under Joint Forest Management increased from 22,017,583 ha to 2,144,000 ha in March 2006 with 106,482 recognized Joint Forest Management committees countrywide [57].
Deforestation is a major environmental challenge which has been persistent from the past, and the situation is more worsened at present. Therefore, there is an urgent need to focus on the mitigative measures in order to prevent the distressing effects of deforestation in the near future. In order to alleviate the problem of deforestation, the strategies should be based on the underlying causes of the same. Also, the strategies for mitigating the problem of deforestation require its effective implementation that needs the recognition of the roles of national, state and municipal governments along with the pro-active role of the civil society and private society [36]. The continuous increase in the human population especially in the developing countries has resulted in enhanced pressure on the forests for human settlements and other land use practices. A reduction in the growth rate of human population plays a crucial role in reducing the practice of deforestation [36]. Alternatives to slash-and-burn agriculture can be adopted as a strategy to mitigate the deforestation by boosting the agricultural and forestry productions for the shifting cultivators ([58, 59]). The sustainable alternatives for slash-and-burn cultivation will assist the poor farmers in leading a better life without destroying additional forests [60]. Public policies and laws with greater security, accessibility to the minimal inputs required to maintain or enhance food production and an opportunity to the cultivators to market their products will aid in the sustainability of shifting cultivation systems [61].
Another strategy that can be applied to control the rate of deforestation is through the adoption of sustainable agroforestry, sustainable logging, agro-pastoral production systems, etc. [62]. The sustainable forest management practices can be promoted only if it is ecologically, economically and socially sustainable [36]. Agroforestry has been considered as one of the methods to curb deforestation which in turn aid in reduction of CO2 emissions and mitigation of climate change effect [63, 64]. The adoption of agroforestry practices has resulted in an increase in the income of agroforestry adopters as compared to that of non-agroforestry adopters and has also contributed towards improving soil fertility, reducing deforestation and conserving soil and water [65]. The provision of protected areas is one of the key steps towards an attempt to reduce deforestation which is generally motivated through biodiversity conservation [34]. Also, the global endeavours to reduce tropical deforestation are dependent heavily on the establishment of protected areas. It has been found that protection reduced deforestation as approximately 10% of the protected forests would have been deforested if they would have not been protected [66]. Tropical protected areas reduced deforestation which was liable for around one-tenth of total anthropogenic carbon emissions, thus playing a significant role in mitigating the effects of climate change and protecting biodiversity and ecosystem services [67]. Similarly, a total loss of 15.4% in the unprotected mangrove cover was compensated by the 15.7% rise in the protected mangrove cover (protected by government as per Ramsar Convention) which resulted in a net increase of 13.3% in mangrove cover across India in the last 25 years [68]. Implementation of forest certification can be among one of the strategies to control deforestation around the world. Forest certification is a process through which the producers identify their products in the marketplace and receive greater market accessibility and higher prices for their products by fulfilling certain stringent sustainable forestry standards [69]. Certification has played an important role in protecting Penten forests from 1986 to 2007, and the certified forests experience 20 times less deforestation than non-certified areas [70]. Thus, certification of forest can play a major role in controlling deforestation since the timber certification was found to be negatively related with deforestation, i.e. the increase in the certification process has resulted in a declination in the deforestation rate [71].
The Government of India has been continuously putting significant efforts in terms of conserving the forest resources. They have not only taken measures to conserve the present forest cover but also have initiated several measures to increase the forests and tree cover of the country. Both the national and state governments are jointly responsible for the sustainable management of the forest resources in India. Various steps, rules and laws have been brought and implemented in order to conserve the forest cover of India. Some of these rules and laws are [72]:
The Indian Forest Act, 1927: The act is an amalgamation of laws relating to forests, the transit of forest produce and the duty leviable on timber and other forest produces. It defines the procedures for declaring an area of a reserved forest, a protected forest or a village forest by the state government. With the amendment in the Act in 2012, it also prohibited the fresh clearances in forests and setting fire in a reserved forest.
Forest Conservation Act, 1980 (with an amendment in 1988): The main purpose of the proposition of this act was to conserve the forests and to look into the matters connected therewith or ancillary or incidental thereto. With the implementation of this act, a prior approval of the Central Government is required for any sort of diversion of forest areas for the non-forestry purposes.
[73]: The establishment of the National Forest Policy was also among one of the steps taken by the Government of India in order to ensure compensatory afforestation, essential environmental safeguards, sustainable utilization, maintenance, restoration and enhancement of forest areas.
Wildlife Protection Act, 1972: The wildlife protection act was enacted basically to protect wild animals, birds and plants and for matters connected therewith or ancillary or incidental thereto with a view to ensure the ecological and environmental security of the country.
The Scheduled Tribes and Other Traditional Forest Dwellers (Recognition of Forest Rights) Act, 2006: The act was framed with the purpose to recognize and vest the forest rights and occupation in forest land in forest dwelling scheduled tribes and other traditional forest dwellers who have been residing in such forests for generations but whose rights could not be recorded. These recognized rights of the forest dwelling scheduled tribes and other traditional forest dwellers include the responsibilities and authority for sustainable use, conservation of biodiversity and maintenance of ecological balance which in turn aid in strengthening the conservation regime of the forests while ensuring livelihood and food security of the forest dwellings scheduled tribes and other traditional forest dwellers.
Apart from these laws, the Government of India has also established Forest Survey of India (FSI), an organization under the Ministry of Environment, Forest and Climate Change, whose primary work is to gather and evaluate the country's forest wealth through a nationwide survey to measure forest areas [74]. This in turn aids in determining the factors and drivers behind the loss or gain in forest cover of any particular region in India. Another council, i.e. the Compensatory Afforestation Fund Management and Planning Authority (CAMPA), was established by the Government of India in 2009 as a National Advisory Council under the chairmanship of the Union Minister of Environment, Forest and Climate Change for the monitoring, technical assistance and evaluation of compensatory afforestation activities. This was particularly meant to promote afforestation and regeneration activities as a way of compensating for forest land diverted to non-forest uses [75]. Certain scheme such as Integrated Forest Protection Scheme (IFPS) was being formulated by the government to save the forests from fire. The scheme was designed by combining the forest fire protection and management technique along with forest conservation. Various other missions and programmes such as the National Mission for a Green India (NMGI) and National Afforestation Programme (NAP) were also being launched by the Government of India where the main aim of NMGI was to improve the quality of five million hectares of degraded forests and to bring another five million hectares of non-forest areas under forest cover through social and farm forestry. On the other hand, the NAP was launched with the objective to develop the forest resources with people's participation, with a focus on improving the livelihood of the forest-fringe communities, especially the poor [75].
The Ministry of Environment, Forest and Climate Change has been optimistic in strengthening the role of women in conservation of forest at local community levels since long. The National Forest Policy [73], for the first time, acknowledged the necessity of including woman members in forestry schemes. The Joint Forest Management Policy of 1990 mandated woman representatives not less than 40% in general body and 50% in executive body of the local forestry institutions like the JFM committee. Later in 2002, the Biodiversity Authority of India reframing the local biodiversity management committee structure mandated the reservation of one-third of its members as women. Thus, this understanding of the role of women in the local-level conservation measures and implementation of related rules has aided in improving the management of forest in rural regions of the country [76].
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Receipt of complaints will be acknowledged in writing and Intech Limited will respond fully to concerns within 15 business days.\n\n
Customers have the right to terminate the contract without giving any reason (written notice of termination). The deadline for said termination is fourteen (14) days from the date of receipt of goods. Returns are at the expense of the Customer and must be made within the fourteen (14) days from the date of the written notice of termination. Intech Limited will process refunds to the Customer without undue delay.\n\n
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