From Big Medical Encyclopedia

BLOOD GROUPS — the normal immunogenetic signs of blood allowing to combine people in certain groups on similarity of antigens of their blood. The last received the name of group antigens (see), or isoantigens. Belonging of the person to this or that G. to. is its individual biol, feature, edges begins to form already in the early period of embryonic development and does not change during all subsequent life. Some group antigens (isoantigens) are not only in uniform elements and a blood plasma, but also in other cells and fabrics, and also in secrets: to saliva, an amniotic fluid, went. - kish. juice, etc. The intraspecific isoantigenic differentiation is inherent not only in People, but also animals at whom the special G. are found to.

Knowledge of G. to. doctrines are the cornerstone about hemotransfusion (see), are widely applied in clinical practice and forensic medicine. The genetics of the person and anthropology cannot do without use of group antigens as genetic markers.

There is big literature on G.'s communication to. with various infectious and noninfectious diseases of the person. However this question is in a stage of studying and accumulation of the facts.

Science about G. to. arose at the end of 19 century as one of sections of the general immunology (see). Therefore it is natural that such categories of immunity as concepts about antigens (see) and antibodies (see), their specificity, completely keep the value and during the studying of an isoantigenic differentiation of a human body.

In erythrocytes, leukocytes, thrombocytes, and also in a blood plasma of people openly there are a lot of tens of isoantigens. The most studied isoantigens of erythrocytes of the person are presented in tab. 1 (about isoantigens of leukocytes, thrombocytes, and also isoantigens of serum proteins — see below).

The stroma of each erythrocyte contains in itself a large number of the isoantigens characterizing intraspecific group-specific signs of an organism of people. Apparently, the true number of antigens on a surface of membranes of erythrocytes of the person considerably exceeds number of already open isoantigens. Existence or absence in erythrocytes of this or that antigen, and also various combinations create them a big variety of the antigenic structures inherent in people. If to take into account even not a full range of the isoantigens discovered in uniform elements and in proteins of a blood plasma, then direct calculation will indicate existence of many thousands immunological of distinguishable combinations.

The isoantigens which are in a genetic linkage are united in the groups which received the name of the AB0 systems, a Rhesus factor, etc.

Blood groups of the AB0 system

Blood groups of the AB0 system are opened in 1900 by K. Landshteyner. Mixing erythrocytes of one persons with normal blood sera of others, he found out that he at one combinations of serums and erythrocytes is observed hemagglutination (see) — it is not at others. On the basis of these factors K. Landshteyner came to conclusion that blood of various people is heterogeneous and can be conditionally divided into three groups which it designated letters A, B and S. Vskore after that Dekastello and Shturli (A. Decastello, A. Sturli, 1902) send people, erythrocytes and which serums differed from erythrocytes and serums of the mentioned three groups. They considered this group as a deviation from Landshteyner's scheme. However Ya. Yansky in 1907 established that this G. to. not an exception of Landshteyner's scheme, but independent group, and, therefore, all people on immunol, to properties of blood are divided into four groups.

Distinctions of agglyutinabelny properties of erythrocytes depend on the certain substances, specific to each group, which are available in them — agglutinogens (see. Agglutination ), which according to E. Dungern and L. Girshfeld (1910) offer designate letters A and B. According to this designation erythrocytes of one persons do not contain agglutinogens A and B (the I group on Yansky, or 0 group), erythrocytes of others contain agglutinogen A (the II blood group), erythrocytes of the third parties contain agglutinogen B (the III blood group), erythrocytes of the fourth contain agglutinogen A and B (the IV blood group).

Depending on existence or absence in erythrocytes of group antigens A and B in plasma there are normal (natural) isoantibodies (Hemagglutinins) in relation to these antigens. At faces of group 0 two types of group antibodies contain: anti-And yes anti-In (an alpha and a beta). At faces of group A the isoantibody r contains (anti-In), persons have groups B — an isoantibody and (anti-And) and at faces of the AV group both hemagglutinins are absent. Ratios between isoantigens and isoantibodies are presented in tab. 2.




Is accepted alphabetic, but not digital designation G. to., and also full writing of a formula to., considering both antigens of erythrocytes, and antibodies of serum (0αβ, Aβ, Bα, AB0). Apparently from tab. 2, the blood group is characterized equally both by isoantigens, and isoantibodies. At G.'s definition to. it is necessary to consider both of these indicators as persons can meet slight isoantigens of erythrocytes and persons at whom isoantibodies are insufficiently active or are even absent.

E. Dungern and Girshfeld (1911) found that group antigen A is not homogeneous and can be subdivided into two subgroups — A1 and A2 (on the terminology offered by K. Landshteyner). Erythrocytes of subgroup of A1 are well agglutinated by the corresponding serums, and erythrocytes of subgroup of A2 — is weak, and it is necessary to apply them to identification highly active standard serums of the Bα group y 0αβ. Erythrocytes of the A1 group meet in 88%, and the A2 groups — in 12%. Further options of erythrocytes with even more poorly expressed agglyutinabelny properties were found: A3, A4, A5, Az, A0, etc. Such slaboagglyutiniruyushchikhsya options of erythrocytes of group A it is necessary to reckon with a possibility of existence in practice of G.'s definition to., in spite of the fact that they meet very seldom. Group

antigen B, unlike antigen A, is characterized by bigger uniformity. Rare options and this antigen — B2, B3, by Bw, Vkh, etc. are described, however. The erythrocytes containing one of these antigens had poorly expressed agglyutinabelny properties. Use of highly active standard Aβ and 0αβ serums allows to reveal also these slight agglutinogens B.

Erythrocytes of group 0 are characterized not only by lack in them of agglutinogens A and B, but also existence of special specific antigens of H and 0. H and 0 antigens contain not only in erythrocytes of group 0, but also in erythrocytes of subgroup of A2 and least of all — in erythrocytes of subgroup of A1 and A1B.

If availability of antigen H in erythrocytes does not raise doubts, then the issue of independence of existence of antigen 0 finally is not resolved yet. According to Morgan and Watkins's researches (W. Morgan, W. Watkins, 1948), distinctiveness of antigen H is existence it in biol, liquids of secretaries of group substances and absence — at nesekretor. Antigen 0, unlike antigen N, A and B, with secrets is not emitted.

The great value in practice of definition of antigens of the AB0 system, and in particular subgroups of A1 and A2, was gained opened by Boyd (W. Boyd, 1947, 1949) and irrespective of it Renkonen (To. Renkonen, 1948) vegetable matters — phytohemagglutinins. Phytohemagglutinins, specific concerning group antigens, are called also lectins (see). «Pectins find in seeds of bean plants this more often. Leguminosa. Water-salt extracts from seeds of Dolichos biflorus and Ulex europeus can serve as a perfect mix of phytohemagglutinins for definition of subgroups in groups A and AB. The lectins received from seeds of Dolichos biflorus react with erythrocytes of the A1 and A1B group and do not react with erythrocytes groups of A2 and A2B. The lectins received from seeds of Ulex europeus, on the contrary, react with erythrocytes of the A2 and A2B group. Lectins from seeds of Lotus tetragonolobus and Ulex europeus apply to detection of antigen N.

Lectins (anti-In) in relation to erythrocytes of group B are found in seeds of Sophora japonica.

The lectins reacting with antigens of other systems G. to are found. Also specific fitopretsipitina are found.

Peculiar antigenic it is gray l, the option of blood was found by Y. Bhende with soavt, in 1952 in the resident of Bombay, erythrocytes to-rogo did not contain any of the known antigens of the AB0 system, and in serum there were antibodies anti-And, anti-In and Anti-N; this option of blood received the name «Bombay» (Oh). Further the option of blood like Bombay was found at people and in other parts of the globe.

Antibodies in relation to group antigens of the AB0 system happen normal, naturally arising in the course of formation of an organism, and immune, shown as a result of immunization of the person, e.g. at administration of inogruppny blood. Normal isoantibodies anti-And yes anti-In are usually immunoglobulins M (IgM) and are more active at lowered (20 — 25 °) temperature. Immune group isoantibodies are more often connected with immunoglobulins G (IgG). In serum all three classes of group immunoglobulins can meet, however, (IgM, IgG and IgA). In milk, saliva, a phlegm often there are antibodies of secretory type (IgA). Apprx. 90% of the immunoglobulins found in colostrum belong to the class IgA. The antiserum capacity of IgA in colostrum is higher, than in serum. At faces of group 0 both types of antibodies (anti-A and anti-B) belong usually to one class immunoglobulins (see). Both IgM, and IgG group antibodies can have hemolitic properties, i.e. connect a complement in the presence in a stroma of erythrocytes of the corresponding antigen. On the contrary, antibodies of secretory type (IgA) of hemolysis do not cause as do not connect a complement. Agglutination of erythrocytes requires 50 — 100 times less molecules IgM of antibodies, than the molecules IgG of group antibodies.

Normal (natural) group antibodies begin to appear at the person in the first months after the birth and reach the maximum caption approximately by 5 — 10 years. After that the antiserum capacity remains at rather high level for many years, and then with age there is its gradual decrease. The caption of hemagglutinins anti-And normal varies within 1: 64 — 1: 512, and a caption of hemagglutinins anti-In — within 1:16 — 1: 64. In rare instances natural Hemagglutinins can be expressed poorly that complicates their identification. Such cases are observed at a hypogammaglobulinemia or agammaglobulinemias (see). In addition to hemagglutinins, in serums of healthy people also normal group hemolysins meet (see. Hemolysis ), but in a low caption. Hemolysins anti-And, as well as corresponding to them agglutinins, are more active, than hemolysins of Anti-century.

The person can have also immune group antibodys as a result of parenteral receipt in an organism of incompatible antigens in the group relation. Such processes of isoimmunization can take place at transfusion both whole incompatible blood, and its separate ingredients: erythrocytes, leukocytes, plasma (serum). Most often immune antibodys anti-And which are formed at persons of a blood group 0 and Century meet. Immune antibodys anti-In meet less often. Introduction to an organism of the substances of animal origin similar to group antigens A and B of the person can lead to emergence of group immune antibodys also. Immune group antibodys can appear also as a result of isoimmunization during pregnancy in case of belonging of a fruit to a blood group incompatible with a blood group of mother. Immune hemolysins and Hemagglutinins can result and parenteral use in to lay down. - professional, the purposes of some drugs (serums, vaccines, etc.) containing substances, similar to group antigens.

Substances, similar to group antigens of the person, are eurysynusic in the nature and can be the cause of immunization. These substances are found also in some bacteria. From this it follows that some infections can also stimulate formation of immune antibodys in relation to erythrocytes of group A and B. Formation of immune antibodys in relation to group antigens is of not only theoretical interest, but has also great practical value. Persons 0αβ reckon with a blood group as usually universal donors, i.e. their blood can be transfused to faces of all groups without exception. However the provision on the universal donor is not absolute as faces of group 0, which hemotransfusion owing to availability in it of immune hemolysins and hemagglutinins a high caption can meet (1: 200 and more) can lead to lethal outcomes. Among universal donors, thus, there can be also «dangerous» donors and therefore blood of these persons can be transfused only to patients with (0) blood group of the same name (see. Hemotransfusion ).

Group antigens of the AB0 system, in addition to erythrocytes, were found also in leukocytes and thrombocytes. I. L. Krichevsky and L. A. Schwarzman (1927) for the first time found group antigens A and B in the fixed cells of various bodies { a brain, a spleen, a liver, a kidney). They showed that bodies of people of a blood group And, as well as their erythrocytes, contain antigen A, and bodies of people of a blood group In respectively erythrocytes possess

antigen B. Further group antigens were found almost in all tissues of the person (muscles, skin, a thyroid gland), and also in cells of benign and malignant tumors of the person. The exception was made by lens, in Krom group antigens are not found. Antigens A and B are found in spermatozoa, liquid of sperm. The amniotic fluid, saliva, a gastric juice are especially rich with group antigens. Few group antigens in blood serum and in urine, and in cerebrospinal liquid they are practically absent.

Sekretora and nesekretor of group substances. On ability to emit group substances with secrets of all people divide into two groups: secretaries (Se) and nesekretor (se). On materials R. M. Urinson (1952), 76% of people are secretaries and 24% — nesekretor of group antigens. Existence of intermediate groups between strong and weak secretaries of group substances is proved. Content of group antigens in erythrocytes of secretaries and nesekretor equally. However in serum and in fabrics of bodies of nesekretor group antigens are found in weaker degree, than in fabrics of secretaries. Ability of an organism to emit group antigens with secrets is descended on dominant type. Children whose parents treat nesekretor of group antigens are also nesekretor. The faces possessing a dominant gene of secretion are capable to excrete the group substances, faces having a recessive gene of not secretion with secrets, have no this ability.

Biochemical nature and properties of group antigens. Group antigens A and B of blood and bodies are steady against effect of alcohol, ether, chloroform, acetone and formalin, high and low temperature. Group antigens A and B in erythrocytes and in secrets are connected with various molecular structures. Group antigens A and B of erythrocytes are glycolipids (see), and group antigens of secrets — glycoproteins (see). The group glycolipids A and B emitted from erythrocytes contain fat to - you, sphingosine and carbohydrates (glucose, a galactose, a glycosamine, galaktozamin, to a fukoz and sialine to - that). A carbohydrate part of a molecule is connected with fat to-tami through sphingosine. The glycolipidic drugs of group antigens emitted from erythrocytes are haptens (see); they specifically react with the corresponding antibodies, but are not capable to cause products of antibodies in immunizirovanny animals. Accession to this hapten of protein (e.g., horse serum) turns group glycolipids into full-fledged antigens. It gives the chance to conclude, as in native erythrocytes which are full-fledged antigens group glycolipids are connected with protein. The purified group antigens emitted from cystous liquid of an ovary contain 85% of carbohydrates and 15% of amino acids. Average pier. the weight of these substances makes 3 X X 105 — 1 x 106 dalton. Aromatic amino acids are present only at very insignificant quantities; the amino acids containing sulfur are not found. Group antigens A and B of erythrocytes (glycolipids) and secrets (glycoproteins), though are connected with various molecular structures, have identical antigenic determinants. Group specificity of glycoproteins and glycolipids is defined by carbohydrate structures. The small number of the sugars which are located on the ends of a carbohydrate chain is an important part of a specific antigenic determinant. As showed the chemical analysis [W. Watkins, 1966], in composition of antigens A, B, Lea enter identical carbohydrate components: alpha hexose, a D-galactose, alpha methyl-pentose, L-fukoza, two aminosugars — N-acetyl a glycosamine and N - acetyl - D - galaktozamin and N-atsetilneyra-minovaya to - that. However forming from these carbohydrates of structure (antigenic determinants) are not identical, as defines specifics of group antigens. L-fukoza plays an important role in structure of a determinant of antigen N, N - acetyl - D - galaktozamin — in structure of a determinant of antigen A, and D - galak-toza — in structure of a determinant of group antigen B. Peptide components do not take part in structure of determinants of group antigens. They as assume, promote only strictly certain arrangement in space and orientation of carbohydrate chains, give them a certain rigidity of structure.

Genetic control of biosynthesis of group antigens. Biosynthesis of group antigens is carried out under control of the corresponding genes. A certain order of sugars in a chain of group polysaccharides is created not by the matrix mechanism as for proteins, and results from strictly coordinate effect of specific glikozil-transferazny enzymes. According to Watkins (1966) hypothesis, group antigens which structural determinants are carbohydrates can be considered as afterproducts of genes. Primary products of genes are proteins — the glikoziltransferaza catalyzing transfer of sugars from a glikozilny derivative nukleoziddifosfat on carbohydrate chains of a glikoproteinapredshestvennik. Serol., genetic and biochemical, researches give the grounds to assume that genes And, In and Le control glikoziltransferazny enzymes which catalyze accession of the corresponding units of sugars to carbohydrate chains of a preformirovanny glycoprotein molecule. Recessive alleles of these loci function as inactive genes. The chemical nature of substance predecessor in a due measure is not defined. One researchers consider that the general for all group antigens predecessors is the glikoproteidny substance identical on the specificity to polysaccharide of a pneumococcus of the XIV type. On the basis of this substance are under construction under the influence of genes And, In, N, Le the corresponding antigenic determinants. Substance of antigen H is the main structure, edges enters all group antigens of the AB0 system. Other researchers [Feyzi, Kabat (T. Feizi, E. Kabat), 1971] produced the evidence that the predecessor of group antigens — substance of antigen I.

Isoantigens and isoantibodies of the AB0 system in ontogenesis. Group antigens of the AB0 system begin to be found in erythrocytes of the person in the early period of its embryonic development. Group antigens found in erythrocytes of a fruit on the second month of embryonal life. Having early created in erythrocytes of a fruit, group antigens A and B reach the greatest activity (sensitivity to the corresponding antibodies) by three years of life. The Agglyutinabelnost of erythrocytes of newborns makes 1/5 part of an agglyutinabelnost of erythrocytes of adults. Having reached a maximum, the caption of agglutinogens of erythrocytes within several decades remains at a fixed level, and then its gradual decrease is observed. Specificity of an individual group differentiation inherent in each person remains during all his life regardless of the postponed infectious and noninfectious diseases, and also from impacts on an organism various physical. - chemical factors. During all individual human life there are only quantitative changes in a caption of its group gemagglyutinogen And yes In, but not qualitative. In addition to age changes about which it was told above a number of researchers noted decrease in an agglyutinabelnost of erythrocytes of group A at patients with a leukosis. Assume that at these persons change of process of synthesis of predecessors of antigens A and B took place.

Inheritance of group antigens. Soon after opening at G.'s people to. it was noted that group antigenic serol. properties of blood of children are in strictly certain dependence on group accessory of blood of their parents. E. Dungern and L. Girshfeld as a result of inspection of families came to conclusion that group signs of blood are descended by means of two genes independent from each other which they designated, as well as the antigens corresponding to them, letters A and V. Bernstein (F. Bernstein, 1924), based on laws of inheritance of G. Mendel, subjected to the mathematical analysis the facts of inheritance of group signs and came to conclusion about existence of the third genetic sign defining group 0. This gene, unlike dominant genes And yes In, is recessive. According to Furukhata's theory (T. Furuhata, 1927), are descended the genes defining development not only antigens A, B and 0(H) but also hemagglutinins an acorus. Agglutinogens and agglutinins are inherited in correlative communication in the form of the following three genetic signs: 0αβр, Aβ and Bα. Antigens A and B are not genes, but develop under specific influence of genes. The blood group, as well as any ancestral feature, develops under specific influence of two genes from which one comes from mother, and another — from the father. If both genes are identical, then an oospore and consequently, and the organism which developed from it will be homozygous; if the genes defining the same sign are not identical, then the organism will have heterozygous properties.

According to it a genetic formula G. to. not always matches phenotypical. E.g., to a phenotype 0 there corresponds the genotype 00, to a phenotype And — a genotype of AA and the joint-stock company, to a phenotype In — a genotype of B B and IN, to a phenotype of AV — a genotype of AV.

Antigens of the AB0 system unequally often occur among various people. Frequency, about G.'s cut to. occur among the population of some cities of the USSR, it is presented on tab. 3.

To. the AB0 systems have paramount value in practice of hemotransfusion, and also at selection of compatible couples of donors and recipients at organ transplantation of fabrics (see. Transplantation ). About biol. value of isoantigens and isoantibodies is known a little. Assume that normal isoantigens and isoantibodies of the AB0 system play a role in maintenance of constancy internal environment of an organism (see). There are hypotheses of protective function of antigens of the AB0 system of a digestive tract, seed and amniotic liquid.

A blood group of the Rh system

Blood groups of the Rh (Rhesus) system take the second place on value for medical practice. This system received the name from monkeys of rhesus whose erythrocytes were applied by K. Landshteyner and A. Winer (1940) to immunization of rabbits and Guinea pigs from whom specific serums were received. By means of these serums in erythrocytes of the person found Rh antigen (see. Rhesus factor ). The greatest progress in studying of this system was made thanks to receiving isoimmune serums from the multigiving birth women. This one of the most complex systems of an isoantigenic differentiation of a human body includes more than twenty isoantigens. In addition to five main R h antigens (D, With, with, E, e), enter this system also their numerous options. One of them are characterized by the lowered agglyutinabelnost, i.e. differ from the main R h antigens in a quantitative sense, other options have qualitative antigenic features.

Achievements of the general immunology are considerably connected with studying of antigens of the Rh system: opening of the blocking and monovalent antibodies, development of new methods of a research (Koombs's reaction, hemagglutination reaction in colloid environments, use of enzymes in immunol, reactions, etc.). Progress in diagnosis and prevention of a hemolitic disease of newborns (see) is also reached by hl. obr. during the studying of this system.

The blood group of the MNSs system

Seemed that the system of group antigens M and N opened by K. Landshteyner and F. Levin in 1927 is rather well studied and consists of two main antigens — M and N (such name is given antigens conditionally). Further researches, however, showed that this system is not less difficult, than the Rh system, and includes apprx. 30 antigens (tab. 1). Antigens M and N were discovered by means of the serums received from rabbits immunizirovanny by erythrocytes of the person. People have antibodies of Anti-m and in particular anti-N meet seldom. On many thousands of transfusions of blood, incompatible concerning these antigens, only isolated cases of formation of isoantibodies of Anti-m or anti-N were celebrated. On the basis of it group accessory of the donor and recipient on the MN system in practice of hemotransfusion is usually not considered. Antigens M and N can be in erythrocytes together (MN) or everyone (M and N). According to A. I Rozanova's (1947) data, inspected edges in Moscow 10 000 people, persons of a blood group of M meet in 36%, groups N — in 16%, and the MN groups — in 48% of cases. By the chemical nature antigens M and N are glycoproteins. Enters into structure of antigenic determinants of these antigens neuraminic to - that. Its eliminating from antigens by processing of the last a neuraminidase of viruses or bacteria leads to an inactivation of antigens M and N.

Formation of antigens M and N happens in the early period of an embryogenesis, antigens are found in erythrocytes of embryos 7 — 8 weeks age. Beginning with the 3rd month antigens M and N in erythrocytes of embryos are well-marked and do not differ from antigens of erythrocytes of adults. Antigens M and N are descended. One sign (Or N) the child receives m from mother, another — from the father. It is established that children can have only those antigens which are available for parents. In the absence of this or that sign at parents children also cannot have them. On the basis of it the MN system matters in court. - medical practice at the solution of questions of a doubtful paternity, motherhood and substitution of children.

In 1947 by means of the serum received from the multigiving birth woman, Walsh and Montgomery (R. Walsh, S. of Montgomery) discovered the antigen S connected with the MN system. A little later also antigen s was found in erythrocytes of the person.

Antigens S and s are controlled by allelic genes (see. Alleles ). At 1% of people antigens S and s can be absent. To. these persons are designated a Su symbol. In addition to MNSs antigens, find the complex antigen U consisting of components of antigens S and s in erythrocytes of some persons. Also other diverse options of antigens of the MNSs system meet. One of them are characterized by the lowered agglyutinabelnost, others — have qualitative antigenic distinctions. In erythrocytes of people also the antigens (Not, etc.) which are genetically connected with the MNSs system were found.

Blood groups of system P

Along with antigens M and N K. Landshteyner and F. Levin (1927) discovered antigen P in erythrocytes of the person. Depending on existence or lack of this antigen all people were divided into two groups — P+ and P —. Long time was considered that the system P is limited to existence only of these two options of erythrocytes, however further researches showed, as this system is more difficult. It turned out that erythrocytes of most of R-negative subjects contain the antigen coded by other allelomorphic gene of this system. This antigen was called P2, unlike P1 antigen which was designated as P+ earlier. There are persons at whom both antigens (P1 and P2) are absent. Erythrocytes of these persons designate a letter of river. Later antigen of Rk was discovered and the genetic linkage of both this antigen, and Tja antigen with system P is proved. Consider [R. Sanger, 1955] that Tja antigen is a complex of P1 and P2 antigens. Faces of the P1 group meet in 79%, the P2 groups — in 21% of cases. Faces of group of Rk and r meet very seldom. Serums for detection of antigens P receive both from people (isoantibody), and from animals (heteroantibody). As from - and heteroantibodies Anti-R belong to the category of vhole antibodies of cold type as the agglutination test caused by them occurs best of all at t ° 4 — 16 °. Antibodies Anti-R, active are described and at the body temperature of the person. Isoantigens and isoantibodies of system P have the defined wedge, value. Cases of early and late abortions which cause isoantibodies of Anti-river were are noted. Several cases of the posttransfusion complications connected with incompatibility of blood of the donor and recipient on system of antigens P are described.

The established connection between system P and a cold paroxysmal haemoglobinuria of Donat — Landshteynera is of great interest (see. Immunohematology ). Origins of autoantibodies in relation to own P1 and P2 antigens of erythrocytes remain still unknown.

Blood groups of the Kell system

Kell Antigen (Kel) was discovered by Koombs, Murant, Flight (R. Coombs, A. Mourant, R. Race, 1946) in erythrocytes of the child having a hemolitic disease. The name is given antigen by the name of families, at members a cut were for the first time found Kell (K) antigen and K. U antibodies of mother were found the antibodies reacting with erythrocytes of her husband, child and 10% of samples of the erythrocytes received from other persons. Transfused to this woman blood from her husband that, apparently, promoted isoimmunization.

On the basis of availability of antigen K in erythrocytes or its absence all people can be divided into two groups: Kell-positive and Kell-negative. In three years after discovery of antigen K it was established that the Kell-negative group is characterized not just by lack of antigen K, and availability of other antigen — to. Allen and Lewis (F. Allen, S. Lewis, 1957) found serums which allowed to discover Kr's antigens and Krv relating to the Kell system in erythrocytes of people. Stroup, Mack-Ilra (M. of Stroup, M. Macllroy) and sotr. (1965) showed that antigens of the Sutter group (Jsa and Jsb) are also genetically connected with this system. Thus, the Kell system, as we know, includes three: vapors of antigens: To, to; Kra; KRD; Jsa and JsB which biosynthesis is coded by three couples of allelic genes To, k; Kpb, Krv; Jsa and Jsb. Antigens of the Kell system are descended under the general genetic laws. Formation of antigens of the Kell system belongs to the early period of an embryogenesis. In erythrocytes of newborns these antigens are rather well-marked. Antigens Kik have rather high immunogene activity. Antibodies to these antigens can arise as in the course of pregnancy (in the absence of this or that antigen at mother and existence them at a fruit), and as a result of repeated hemotransfusions, incompatible concerning Kell antigens. Many cases of hemotransfusionic complications and a hemolitic disease of newborns whose cause isoimmunization antigen K was are described. Antigen K, according to T. M. Piskunova (1970), edge inspected 1258 residents of Moscow, was at 8,03% and was absent (kk group) at 91,97% inspected.

Katbush, Mollison and Parkin (M. of Cutbush, P. Mollison, D. Parkin, 1950) send blood groups of the Duffy system at the patient with hemophilia of an antibody which reacted with unknown antigen. The last was: the patient, or in abbreviated form Fya is called by them Duffy antigen (Daffi), by last name. Soon after that also the second antigen of this system — Fyb was found in erythrocytes. Antibodies in relation to these antigens receive or from patients, to - an eye repeated hemotransfusions, or from women whose newborn children had a hemolitic disease were made. Full meet and monovalent antibodies are more often and therefore it is necessary to apply Koombs's reaction to their detection (see. Koombs reaction ) or to put an agglutination test in the colloid environment. To. Fy (a+b —) meets in 17,2%, the Fy group (and — b+) — in 34,3% and the Fy group (a+b+) — in 48,5%. Fya and Fyb antigens are descended as dominant characters. Formation of Fy antigens happens in the early period of an embryogenesis. Fya antigen can entail heavy posttransfusion complications at hemotransfusion if not to consider incompatibility to this antigen. Fyb antigen, unlike Fya antigen, is less isoantigenic. Antibodies in relation to it meet less often. Fya antigen is of great interest to anthropologists as it occurs at one people rather often, and at others is absent.

Blood groups of the Kidd system

opened Antibodies to antigens of the Kidd system (Kidd) in 1951. Allen, Daymond and Nedzelya (F. Allen, L. Diamond, V. of Niedziela) at the woman by the name of Kidd, the newborn child a cut had a hemolitic disease. The corresponding antigen in erythrocytes was designated by the letters Jka. Soon after that the second antigen of this system — Jkb was found. Jka and Jkb antigens are a product of function of allelic genes. Jka and Jkb antigens are descended under the general laws of genetics. It is established that children can have no antigens which are absent at their parents. Jka and Jkb antigens occur at the population approximately equally often — in 25%, 50% of people in erythrocytes have both antigens. Antigens and antibodies of the Kidd system have a certain practical value. They can be the cause of a hemolitic disease of newborn and posttransfusion complications at repeated transfusion of this system of blood, incompatible on antigens.

Blood groups of the Lewis system

the First antigen of the Lewis system (Lewis) A. Mourant in 1946 in erythrocytes of the person by means of the serum received from the woman by the name of Lewis was open. This antigen was designated by the letters Lea. In two years Andresen (P. Andresen, 1948) reported about discovery of the second antigen of this system — Leb. M. I. Potapov (1970) found new antigen of the Lewis — Led system on a surface of erythrocytes of the person that expanded our ideas of system of Lewis isoantigens and gave the grounds to assume about existence of an allele of this sign — Lec. Thus, the following G.' existence is possible to. Lewis systems: Lea, Leb, Lec, Led. Antibodies of anti-Le of hl. obr. natural origin. However the antibodies which resulted and immunizations, napr, in the course of pregnancy meet, but it is observed seldom. Agglutinins anti-Le belong to antibodies of cold type, i.e. they are more active at low (16 °) temperature. In addition to serums of a human origin, also immune serums were received from rabbits, goats, hens. Grubb (R. Grubb, 1948) established dependence between Le antigens and ability of an organism to emit group substances AVN with secrets. Leb and Led antigens occur at secretaries of group substances AVN, and Lea and Lec antigens — at nesekretor. In addition to erythrocytes, antigens of the Lewis system are found in saliva and in blood serum. Flight and other researchers consider that antigens of the Lewis system are primary antigens of saliva and serum and only for the second time they prove as antigens on a surface of a stroma of erythrocytes. Le antigens are descended. Formation of Le antigens is defined not only genes of Le, but also is under direct influence of genes of secretion (Se) and not secretion (se). Antigens of the Lewis system unequally often occur at the different people and as genetic markers are of undoubted interest to anthropologists. Exceptional cases of the posttransfusion reactions caused by antibodies of anti-Lea are described and are even more rare — antibodies of anti-Leb.

Blood groups of the Lutheran system

discovered the First antigen of this system S. Callender and R. Race in 1946 by means of the antibodies received from the patient, blood repeatedly was transfused to Krom. Antigen was called by last name sick Lutheran (Lutherans) and designated by the letters Lua. In several years also the second antigen of this system — Lub was discovered. Lua and Lub antigens can meet separately and together with the following frequency: Lua — in 0,1%, Lub — in 92,4%, Lua, Lub — in 7,5%. Agglutinins anti-Lu are more often than cold type, i.e. the optimum of their reaction lies not above t ° 16 °. Very seldom antibodies of anti-Lub and even less often than anti-Lua can be the cause of posttransfusion reactions. There are messages on value of these antibodies in an origin of a hemolitic disease of newborns. Lu antigens are defined already in erythrocytes of umbilical blood. A wedge, value of antigens of the Lutheran system in comparison with other systems is rather small.

Blood groups of the Diego system

opened Diego Isoantigen (Diego) in 1955 for Leyriss, Rent, Sisko (M. of Layrisse, T. Arends, R. Sisco) in erythrocytes of the person by means of the monovalent antibodies found in mother the newborn child a cut had a hemolitic disease. On the basis of existence or lack of Diego (Dia) antigen Indians of Venezuela could be divided into two groups: Di (and +) and Di (and —). In 1967 Thompson, Childere and Hetcher (R. of Thompson, D. Childers, D. Hatcher) reported about stay by them at two Mexican Indians of antibodies of anti-Dih, i.e. the second antigen of this system was discovered. Antibodies of anti-Di — an incomplete form and therefore for G.'s definition to. Diego apply Koombs's reaction. Diego antigens are descended as dominant characters, by the time of the birth are well developed. On materials, accurate O. Prokop, G. Uhlenbruck in 1966, Dia antigen was found in residents of Venezuela (different tribes), Chinese, Japanese, but it was not found at Europeans, Americans (white), Eskimos (Canada), Australians, Papuans and Indonesians. Unequal frequency about what Diego antigen it is widespread among various people, is of great interest to anthropologists. Consider that Diego antigens are inherent in the people of the Mongolian race.

Blood groups of the Auberger system

Au Isoantigen was discovered thanks to joint efforts of fr. and English scientists [Salmon, Liberzh, Sanger (S. by Salmon, G. Liberge, R. Sanger), etc.] in 1961. The name is given this antigen on the first letters of a surname of Auberger (Oberzhe) — women, at a cut antibodies were found. Monovalent antibodies arose, apparently, as a result of repeated hemotransfusion. Au antigen is found at 81,9% of the inspected residents of Paris and London. It is descended. In blood of newborns Au antigen is well-marked.

Blood groups of the Dombrock system

Do Isoantigen were opened by J. Swanson with soavt, in 1965 by means of the monovalent antibodies received from the woman by the name of Dombrock (Dombrok), edges there was an immunizirovana as a result of hemotransfusion. On materials of inspection of 755 inhabitants of Northern Europe (Sanger, 1970), this antigen occurred at 66,36% — the Do group (and +) and was absent at 33,64% — the Do group (and —). Doa antigen is descended as a dominant character; in erythrocytes of newborns this antigen is well-marked.

Blood groups of the Ii system

In addition to the group signs of blood described above, were found in erythrocytes of people also isoantigens from which one are very eurysynusic, and others, on the contrary, occur very seldom (e.g., at members of one family) and approach individual antigens. From eurysynusic antigens G. have the greatest value to. Ii systems. A. Winer, Unger * Coen, Feldman (L. Unger, S. Cohen, J. Feldman, 1956) received from the person who had the acquired hemolitic anemia, antibodies of cold type by means of which it was succeeded to find the antigen designated by the letter «I» in erythrocytes of people. From 22 000 inspected samples of erythrocytes only 5 did not contain this antigen or had it in it is insignificant trace amount. Lack of this antigen was designated the letter «i». Further researches, however, showed that antigen i really exists. Faces of group i have antibodies of anti-I that testifies to qualitative distinction between antigens I and i. Antigens of the Ii system are descended. Antibodies of anti-I are defined in the salt environment as agglutinins of cold type. At the persons having the acquired hemolitic anemia of cold type find usually autoantibodies of anti-I and anti-i. Origins of these autoantibodies remain still unknown. Autoantibodies of anti-i meet at patients with nek-ry forms of a reticulosis, myeloid leukemia, an infectious mononucleosis more often. Antibodies of anti-I of cold type of agglutination of erythrocytes at t ° 37 ° do not give, however they can sensibilize erythrocytes and promote accession of a complement, as leads to a lysis of erythrocytes.

Blood groups of the Yt system

Eton and Moreton (V. of Eaton, J. Morton) with sotr. (1956) found in the person, repeatedly transfused to Krom the blood, antibodies capable to reveal very eurysynusic Yta antigen. Later also the second antigen of this system — Ytb was discovered. Yta antigen — one of the most eurysynusic. It occurs at 99,8% of people. Ytb antigen meets in 8,1% of cases. Distinguish three phenotypes of this system: Yt (a + b-), Yt (and + b +) and Yt (and — b +). Persons of a phenotype Y t (and — b —) are not found. Yta and Ytb antigens are descended as dominant characters.

Blood groups of the Xg system

All group isoantigens about which still there was a speech do not depend on a floor. They with an identical frequency occur both at men, and at women. However Mann (J. Mann) and sotr. in 1962 established that there are group antigens which hereditary transfer happens through a gonosome of X. Newly opened antigen in erythrocytes of people was designated by Xg. Antibodies to this antigen were found at the patient who had a family teleangiectasia. On the occasion of profuse nasal bleedings repeatedly transfused to this patient blood, as was, apparently, the reason of his isoimmunization. Depending on existence or absence in erythrocytes of Xg antigen all people can be divided into two groups: Xg(a+) and Xg (a —). At men Xg(a+) antigen occurs in 62,9% of cases, and at women — in 89,4%. It was established that if both parents treat the Xg group (a —), then and at their children — both boys, and girls — this antigen does not contain. If the father of the Xg(a+) group, and mother of the Xg group (a —), all boys have the Xg group (a —) as in these cases spermatozoa only with a chromosome of Y defining a male of the child come to an ovum. Xg antigen is a dominant character, at newborns it is well developed. Thanks to group Xg antigen the possibility of the solution of a question of an origin of some diseases connected with a floor opened (defects of formation of some enzymes, diseases with Klaynfelter's syndromes, Turner, etc.).

Seldom found blood groups

Along with eurysynusic are described also quite seldom found antigens. E.g., Bua antigen is found by Anderson (S. Anderson) with sotr. in 1963 at 1 of 1000 inspected, and antigen of Vkh — W. Jenkins with sotr. in 1961 at 1 of 3000 inspected. Also even more seldom antigens which are found in erythrocytes of the person are described.

A technique of definition of blood groups

the Technique of definition of blood groups — identification in erythrocytes of group antigens by means of standard serums, and for groups of the AB0 system as well identification of agglutinins in serum of the studied blood by means of standard erythrocytes.

For definition of any one group antigen serums of one specificity are used. Simultaneous use of serums of different specificity of the same system gives the chance to determine full group accessory of erythrocytes by this system. E.g., in the Kell system use only of serum anti-To or only anti-k gives the chance to establish whether they contain the studied erythrocytes a factor To or to. Use of both of these serums allows to resolve an issue of belonging of the studied erythrocytes to one of three groups of this system: KK, Kk, kk.

Standard serums for G.'s definition to. prepare from blood of people, antibody-containing — normal (AB0 systems) or isoimmune (the systems Rh, Kell, Duffy, Kidd, Lutheran, antigens S and s). For definition of group M, N, P and Le antigens most often receive heteroimmune serums.

Technology of definition depends on character of the antibodies which are contained in serum which happen full (normal serums of the AB0 system and heteroimmune) or incomplete (the vast majority isoimmune) and show the activity in different environments and at a different temperature what need of use of the different technology of reaction depends on. The method of use of each serum is specified in the accompanying instruction. The net result of reaction during the use of any equipment comes to light in the form of existence or lack of agglutination of erythrocytes. During the definition of any antigen surely join in reaction positive and negative control.

Definition of blood groups of the AB0 system

Necessary reactants: a) standard serums of groups 0αβ (I), Aβ (II), Bα(III) containing active agglutinins, and the AB (IV) groups — control; b) the standard erythrocytes of the A (II) groups and B (III) having well-marked agglyutinabelny properties and groups 0(1) — control.

G.'s definition to. the AB0 systems it is made by an agglutination test at the room temperature on porcelain or any other white plate with a wetted surface.

For G.'s definition to. the AB0 systems there are two ways. 1. By means of the standard serums allowing to establish what group agglutinogens (And or In) are in erythrocytes of the studied blood and on the basis of it to make the conclusion about its group accessory. 2. At the same time by means of standard serums and erythrocytes — a cross way. At the same time existence or lack of group agglutinogens is also defined and, besides, existence or lack of group agglutinins is established (and, 3) that as a result gives the total group characteristic of the studied blood.

At G.'s definition to. the AB0 systems at ill and other persons, the Crimea is supposed to make hemotransfusion, the first way suffices. In special cases, napr, at difficulty in interpretation of result, and also at blood typing of AB0 at donors, use the second way.

At G.'s definition to. in both the first and second way it is necessary to apply two samples (two different series) of standard serum of each group that is one of the measures warning mistakes.

At the first way blood can be taken from a finger, a lobe of an ear or a heel (from babies) just before definition. At the second (cross) way blood is taken previously from a finger or a vein in a test tube and investigated after coagulation, i.e. after division into serum and erythrocytes.

Fig. 1. Blood typing by means of standard serums. On a plate at previously written designations 0αβ (I), Aβ (II) and Bα (III) nakapyvatsya on 0,1 ml of standard serum of each sample. The small drops of blood put with a row carefully mix up with serum. After that plates shake and observe existence of agglutination (positive reaction) or absence it (negative reaction). When agglutination occurred in all drops, do a control research, mixing the studied blood with serum of the AB (IV) group which does not contain agglutinins and shall not cause agglutinations of erythrocytes.

First way (tsvetn. fig. 1). At previously written designations apply 0,1 ml on a plate (on one big drop) standard serum of each sample so that two rows of drops in the following order across from left to right are formed: 0αβ (I), Aβ (II) and Bα (III).

The studied blood is applied by means of a pipette or the end of a glass rod on small (approximately by 10 times smaller) to a drop near each drop of serum.

Blood is carefully mixed with serum dry glass (or plastic) a stick then the plate is periodically shaken, at the same time watching result which is expressed available to agglutination (popozhitelny reaction) or absence it (negative reaction) in each drop. Time of observation of 5 min. For an exception of not specificity of result on a measure of approach of agglutination, but not earlier than in 3 min., in each drop, in a cut there occurred agglutination, add one drop of isotonic solution of sodium chloride and continue observations, shaking a plate within 5 min. When agglutination occurred in all drops, do still a control research, mixing the studied blood with serum of the AB (IV) group, edge does not contain agglutinins and shall not cause agglutinations of erythrocytes.

Interpretation of result. 1. If in one of drops there was no agglutination, it means that the studied blood does not contain group agglutinogens, i.e. belongs to the O (I) group. 2. If serum of group 0 are (I) and In and (III) caused agglutination of erythrocytes, and serum of the Are (II) group yielded a negative take, it means that the studied blood contains agglutinogen A, i.e. belongs to the A (II) group. 3. If serum of group 0αβ (I) and Aβ (II) caused agglutination of erythrocytes, and serum of the Bα (III) group yielded a negative take, it means that the studied blood contains agglutinogen B, i.e. belongs to the B (III) group. 4. If serum of all three groups caused agglutination of erythrocytes, but in a control drop with serum of the AB0 (IV) group reaction negative, it means that the studied blood contains both agglutinogens — And yes In, i.e. belongs to the AB (IV) group.

Fig. 2. Definition of blood groups in the cross way — at the same time by means of standard serums and by means of standard erythrocytes: on a plate at previously inscribed designations (see fig. 1) nakapyvat two rows of standard serums and near each drop — the studied blood. On the bottom of a plate nakapyvat three big drops (in three points), and near them small droplets of standard erythrocytes in the order specified in the drawing — erythrocytes of group 0(I) — are control since they shall not be agglutinated by any serum. Serum with erythrocytes carefully mixes up. After that plates shake and watch result (a detailed explanation in the text of article).

Second (cross) way (tsvetn. fig. 2). At previously inscribed designations, just as at the first way, apply two rows of standard serums of group 0αβ (I), Aβ (II), Bα(III) on a plate and near each drop — the studied blood (erythrocytes). Besides, apply on the bottom of a plate in three points on one big drop of serum of the studied blood, and near them — on one small (approximately by 40 times smaller) to a drop of standard erythrocytes in the following order from left to right: group 0(I), A (II) and V (III). Erythrocytes of group 0(I) are control since they shall not be agglutinated by any serum.

In all drops serum is carefully stirred with erythrocytes and then observe result during the rocking of a plate within 5 min.

Interpretation of result. At a cross way at first the result which turned out in drops with standard serum (two upper rows) is estimated, just as it becomes at the first way. Then the result received in the lower row, i.e. in those drops in which the studied serum is mixed with standard erythrocytes and is estimated, therefore, in it antibodies are defined. 1. If reaction with standard serums indicates belonging of blood to group 0 (I), and serum of the studied blood agglutinates erythrocytes of the A (II) group and B (III) at negative reaction with erythrocytes of group 0 (I), it indicates existence in the studied blood of agglutinins and and 3, i.e. confirms accessory it to group 0αβ(I). 2. If reaction with standard serums indicates belonging of blood to the A (II) group, serum of ispytuyemy blood agglutinates erythrocytes of the B (III) group at negative reaction with erythrocytes of group 0 (I) and A (II); it indicates existence in the studied blood of agglutinin 3» i.e. confirms accessory it to the group A 3 (1G). 3. If reaction with standard serums indicates belonging of blood to the B (III) group, and serum of the studied blood agglutinates erythrocytes of the A (II) group at negative reaction with erythrocytes of group 0 (I) and B (III), it indicates existence in the studied blood of agglutinin and, i.e. confirms accessory it to the Bα (III) group. 4. If reaction with standard serums indicates belonging of blood to the AB (IV) group, and serum yields a negative take with standard erythrocytes of all three groups, it indicates lack of group agglutinins in the studied blood, i.e. confirms accessory it to the AB0 (IV) group.

Definition of blood groups of the MNSs system

Definition of antigens M and N is made by heteroimmune serums, as well as blood groups of the AB0 system, i.e. on a white plate at the room temperature. For a research of two other antigens of this system (S and s) use the isoimmune serums yielding the most accurate result in to indirect test of Koombs (see. Koombs reaction ). Sometimes serums of anti-S contain vhole antibodies, in these cases the research is recommended to be conducted in the salt environment, similar to definition of a Rhesus factor. Comparison of results of definition of all four factors of the MNSs system gives the chance to establish accessory of the studied erythrocytes and one of 9 groups of this system: MNSS, MNSs, MNss, MMSS, MMSs, MMss, NNSS, NNSs, NNss.

Definition of blood groups of the systems Kell, Duffy, Kidd, Lutheran

Definition of these blood groups is made by indirect test of Koombs. Sometimes high activity of antiserums allows to use for this purpose reaction of conglutination using gelatin similar to definition of a Rhesus factor (see. Conglutination ).

Factors of system P and Lewis define definition of blood groups of systems P and Lewis in the salt environment in test tubes or on the plane, and pretreatment of the studied erythrocytes by proteolytic enzyme is applied to more accurate identification of antigens of the Lewis system (papain, trypsin, protelin).

Definition of a Rhesus factor

Definition of the Rhesus factor which is important along with groups the AB0 system most for a wedge, medicine is made in various ways depending on character of antibodies in standard serum (see. Rhesus factor ).

Leukocytic groups

Leukocytic groups — division of people into the groups caused by existence in leukocytes of the antigens independent of antigens of the system AB0, Rh, etc.

Leukocytes of the person have a complex antigenic structure. They contain antigens of the AB0 and MN system, unambiguous with those which are in erythrocytes of the same individual. This situation is based on the expressed ability of leukocytes to cause antibody formation of the corresponding specificity, to be agglutinated by the group isohemagglutinating serums with a high antiserum capacity, and also to specifically adsorb immune antibodys of Anti-m and anti-N. Factors of the Rh system and other antigens of erythrocytes are less expressed in leukocytes.

In addition to the specified antigenic differentiation of leukocytes, special leukocytic groups are allocated.

For the first time the information about leukocytic groups was received by the fr. researcher Zh. Dosse (1954). By means of the immune serum received from persons, the Crimea was made by repeated repeated hemotransfusions, and containing antileukocytic antibodies of the agglutinating character (leukoagglutinating antibodies), the antigen of leukocytes which is found at 50% of the Central European population was revealed. This antigen was included into literature under the name of IAC. In 1959. J. Rood and soavt, added ideas of leukocytic antigens. On the basis of the analysis of results of a research of 60 immune serums with leukocytes of 100 donors authors came to conclusion about existence of other antigens of the leukocytes designated 2,3, and also 4a, 4b; 5a, 5b; 6a, 6b. In 1964 R. Payne with soavt, established LA1 and LA2 antigens.

Contain more than 40 antigens of leukocytes which can be referred to one of three conditionally marked out categories: 1) antigens of the main locus, or general antigens of leukocytes; 2) antigens of granulocytes; 3) antigens of lymphocytes.

The most extensive group is represented by antigens of the main locus (HLA system). They are the general for polymorphonuclear leukocytes, lymphocytes, and also thrombocytes. According to WHO recommendations, use the alphanumeric designation HLA (Human Leucocyte Antigen) for antigens which existence is confirmed in a number of laboratories at parallel researches. Concerning recently open antigens which existence needs further confirmation use designation by a letter w, to-ruyu insert between alphabetic reference of a locus and digital — an allele.

The HLA system — the most difficult of all known systems of antigens. Genetically H LA antigens belong to four subloci (A, B,C, D), each of which combines allelic antigens (see. Immunogenetics ). Subloci are the most studied And yes

V. K to the first sublocus belong: HLA-A1, HLA-A2, HLA-A3, HLA-A9, HLA-A10, HLA-A11, HLA-A28, HLA-A29; HLA-Aw23, HLA-Aw24, HLA-Aw25, HLA-Aw26, HLA-Aw30 „HLA-Aw31, HLA-Aw32, HLA-Aw33, HLA-Aw34, HLA-Aw36, HLA-Aw43a.

The second sublocus possesses antigens: HLA-B5, HLA-B7, HLA-B8, HLA-B12, HLA-B13, HLA-B14, HLA-B18, HLA-B27; HLA-Bw15, HLA-Bw16, HLA-Bw17, HLA-Bw21, HLA-Bw22, HLA-Bw35, HLA-Bw37, HLA-Bw38, HLA-Bw39, HLA-Bw40, HLA-Bw41, HLA-Bw42a.

Rank antigens HLA-Cw1, HLA-Cw2, HLA-Cw3, HLA-Cw4, HLA-Cw5 as the third sublocus.

The fourth sublocus includes antigens HLA-Dw1, HLA-Dw2, HLA-Dw3, HLA-Dw4, HLA-Dw5, HLA-Dw6. The last two subloci are insufficiently studied.

Apparently, not all HLA antigens even of the first two subloci (And yes In) are known since the sum of gene frequencies on each sublocus did not approach unit yet.

Division of the HLA system into subloci represents big progress in area of studying of genetics of these antigens. The system of HLA antigens is controlled by the genes located on C6 to a chromosome on one in a sublocus. Each gene controls synthesis of one antigen. Having a diploid set of chromosomes (see. Chromosomal complement ), theoretically each individual shall have 8 antigens, practically at fabric typing define four HLA antigens of two subloci so far — And yes V. Fenotipicheski can meet several combinations of HLA antigens. It is possible to carry cases when allelic antigens are ambiguous within the first and second sou of loci to the first option. The person is heterozygous on antigens of both subloci. Phenotypical four antigens — two antigens of the first sublocus and two antigens of the second sublocus are found in it.

The second option represents a situation when the person is homozygous on antigens of the first or second sublocus. Such person contains identical antigens of the first or second sublocus. Phenotypical only three antigens are found in it: one antigen of the first sublocus and two antigens of the second sublocus or, on the contrary, one antigen of the second sublocus and two antigens — the first.

The third option covers a case when the person gomozigoten on both subloci. In this case only two antigens, on one each sublocus phenotypical are defined.

The most frequent — the first option genotype (see). Less often in population the second option of a genotype meets. The third option of a genotype is extremely rare.

The division of HLA antigens on subloci allows to predict optional versions of inheritance of these antigens from parents to children.

The genotype H LA antigens of children is defined by ran lotipy, i.e. the linked antigens controlled by the genes located on one chromosome to-ruyu they receive from each of parents. Therefore a half of HLA antigens at the child is always identical with each of parents.

Considering told, it is easy to present four optional versions of inheritance of antigens of leukocytes of the HLA system of subloci And yes V. Teoreticheski coincidence of HLA - anti-genes among brothers and sisters in a family makes 25%.

The important indicator characterizing each HLA-siste-we antigen is not only its arrangement on a chromosome, but also the frequency of its occurrence in populations, or the population distribution having racial features. Frequency of occurrence of antigen is defined by gene frequency, edges represents the part from total number of the studied individuals expressed in shares of unit about a cut each antigen meets. Gene frequency of antigens H of LA system is a constant for a certain ethnic group of the population. According to Zh. Dosse et al., gene frequency for the fr. population makes: HLA-A1 — 0,141, HLA-A2 — 0,256, HLA-A3 — 0,131, HLA-A9 — 0,247, HLA-B5 — 0,143, HLA-B7 — 0,224, HLA-B8 — 0,156. Similar indicators of gene frequencies H LA antigens are established to Yu. M. Zaretskaya and V. S. Fedrunova (1971) for the Russian population. By means of posemeyny researches of various population groups of the globe it was succeeded to establish distinction in the frequency of the found haplotypes. Features in the frequency of HLA haplotypes are explained by distinction of population distribution of antigens of this system at various races.

The great value for applied and theoretical medicine represents determination of quantity of possible HLA haplotypes and phenotypes in the mixed population of people. The number of possible haplotypes depends on amount of antigens in each sublocus and is equal to their work: number of antigens of the first sublocus (A) X of antigens of the second sublocus (V) = quantity of haplotypes, or 19 X 20 = 380.

Calculations indicate that among about 400 people it is possible to find only two people having similarity on two H LA antigens of subloci And yes

of possible combinations of the antigens defining a phenotype find V. Chislo separately for each sublocus. Calculation is made on a formula for definition of number of a combination on two (for heterozygous individuals) and on one (for homozygous individuals) in a sublocus [Mentsel and Richter (G. Menzel, To. Richter), n(n+1)/2, where n — number of antigens in a sublocus.

For the first sublocus the number of antigens is equal 19, for the second — 20.

Number of possible combinations of antigens in the first sublocus — 190; in the second — 210. The number of possible phenotypes for antigens of the first and second sublocus is equal 190 X 210 == 39 900. I.e. on 40 000 approximately only in one case it is possible to meet two unrelated people with an identical phenotype on H LA antigens of the first and second subloci. The quantity H LA phenotypes considerably will increase when the number of antigens in a sublocus With and a sublocus of D is known.

HLA antigens are a comprehensive system. They are found, in addition to leukocytes and thrombocytes, also in cells of various bodies and fabrics (skin, a liver, kidneys, a spleen, muscles, etc.).

Identification of the majority of antigens of the HLA system (loci And, In, C) it is made with the help serol, reactions: limfotsitotoksichesky test, RSK concerning lymphocytes or thrombocytes (see. Reaction of binding complement ). Immune serums, preferential limfotsitotoksichesky character, receive from persons, sensibilized during repeated pregnancies, transplantation of allogenic fabric or by artificial immunization as a result of repeated injections of leukocytes with the known HLA-fenoti-room. Identification H LA antigens of a locus of D is made by means of the mixed culture of lymphocytes.

The HLA system is of great importance in a wedge, medicine and especially at allogenic transplantation of fabrics as discrepancy of the donor and recipient on these antigens is followed by development of reaction of a tissue incompatibility (see. Incompatibility immunological ). In this regard implementation of fabric typing at selection for transplantation of the donor with similar H LA phenotype is represented quite justified.

Besides, distinction of mother and a fruit on antigens H of LA system at repeated pregnancies causes formation of antileukocytic antibodies which can lead to an abortion or death of a fruit.

HLA antigens matter also at hemotransfusion, in particular leukocytes and thrombocytes.

Other system of antigens of leukocytes independent of HLA, antigens of granulocytes are. This system of antigens is tkanespetsifichesky. It is characteristic of cells of a myeloid row. Antigens of granulocytes are found in polymorphonuclear leukocytes, and also cells of marrow; they are absent in erythrocytes, lymphocytes and thrombocytes.

Three granulotsitarny antigens are known: NA-1, NA-2, NB-1.

Identification of system of granulotsitarny antigens is carried out by means of isoimmune serums of the agglutinating character which can be received from repeatedly pregnant women or persons which were exposed to repeated hemotransfusions.

It is established that antibodies against antigens of granulocytes matter at pregnancy, causing short-term neutropenias of newborns. Antigens of granulocytes play also important role in development of not hemolitic transfusion reactions.

The third category of antigens of leukocytes is made by the lymphocytic antigens inherent only in cells of an adenoid tissue. One antigen from this category which received the designation LyD1 is known. It occurs at people with a frequency apprx. 36%. Identification of antigen is made by means of RSK by the immune serums received from the sensibilized persons which were exposed to repeated hemotransfusions or having repeated pregnancies. Value of this category of antigens in transfusiology and transplantology remains poorly studied.

Groups of serum proteins

Serum proteins have group differentiation. Group properties of many serum proteins of blood are open. The research of group of serum proteins finds broad application in forensic medicine, anthropology and, according to many researchers, matters for hemotransfusion. Groups of serum proteins are independent from serol, systems of erythrocytes and leukocytes, they are not connected with a sex, age and are descended that allows to use them in court. - medical practice.

Groups of the following serum proteins are known: albumine, post-albumine, alfa1-globulin (alfa1-antitrypsin), alfa2-globulin, beta1-globulin, lipoproteid, immunoglobulin. Most of groups of serum proteins comes to light by means of an electrophoresis in the hydrolyzed starch, polyacrylamide gel, an agar or on acetate-cellulose, the group of alfa2-globulin (Gc) is defined by method immunoelectrophoresis (see), lipoproteids — method of precipitation in an agar; group specificity of the proteins relating to immunoglobulins is defined immunol, method — reaction of a delay of agglutination by means of auxiliary system: Rh-positive erythrocytes, sensibilized serums an anti-Rhesus factor with the monovalent antibodies containing this or that group antigen of the Gm system.

Immunoglobulins. Among groups of serum proteins genetic heterogeneity has the greatest value immunoglobulins (see), connected with existence of the inherited options of these proteins — the so-called allotypes differing on antigenic properties. It is most important in practice of hemotransfusion, forensic medicine, etc.

Two main systems of allovariants of immunoglobulins are known: Gm and Inv. Characteristic signs of an antigenic structure of IgG are defined by the Gm system (the antigenic determinants which are localized in a S-trailer half of heavy gamma chains). The second system of Inv immunoglobulins is caused by antigenic determinants of light chains and therefore characterizes all classes of immunoglobulins. Antigens of the Gm system and the Inv system determine by method of a delay of agglutination.

The Gm system contains more than 20 antigens (allotypes) which are designated by figures — Gm(1), Gm(2) etc., or letters — Gm (a), Gm(x) etc. The Inv system has three antigens — Inv(1), Inv(2), Inv(3).

Lack of this or that antigen is designated by the sign «—» [e.g., Gm(1, 2-, 4)].

Antigens of immunoglobin systems at persons of various nationalities meet unequal frequency. Gm (1) antigen occurs among the Russian population in 39,72% of cases (M. A. Umnova, etc., 1963). At many nationalities inhabiting Africa, this antigen contains in 100% of cases.

Studying of allovariants of immunoglobulins is important for clinic, genetics, anthropology and is widely used for interpretation of structure of immunoglobulins. In cases agammaglobulinemias (see), as a rule, antigens of the Gm system do not open.

At the pathology which is followed by deep proteinaceous shifts in blood such combinations of antigens of the Gm system which are absent at healthy faces meet. Some patol, changes of blood proteins can mask antigens of the Gm system as if.

Albumine (Al). Polymorphism of albumine occurs at adults extremely seldom. The double strip of albumine — the albumine having bigger mobility at an electrophoresis (AlF) and slower mobility (Als) is noted. See also Albumine .

Post-albumine (Ra). Distinguish three groups: Ra 1-1, Ra 2-1 and Ra 2-2.

alfa1-Globuliny. In the field of alfa1-globulins the big polymorphism of alfa1-antitrypsin (alfa1-AT-globulin) which received designation of the Pi system (protease-inhibitor) is noted. 17 phenotypes of this system are revealed: PiF, PiJ, PiM, Pip, Pis, Piv, Piw, Pix, Piz, etc.

Under certain conditions an electrophoresis alfa1-globulins have big electrophoretic mobility and are located on an elektroforegramma ahead of albumine therefore some authors call them prealbumins.

alfag-Antitripsin treats glycoproteins. It inhibits activity of trypsin and other proteolytic enzymes. Fiziol, a role of alfa1-antitrypsin is not established, however increase in its level at some fiziol, states is noted and patol, processes, napr, at pregnancy, after reception of contraceptives, at an inflammation. Low concentration of alfa1-antitrypsin is connected with an allele of Piz and Pis. Note communication of insufficiency of alfa1-antitrypsin with hron, obturatsionny pulmonary diseases. These diseases people, homozygous long for an allele of Pi2 or Pi2, heterozygous on alleles, and Pis more often.

Connect with insufficiency of alfa1-antitrypsin also the special form of emphysema of lungs which is descended.

α2-Глобулины. In this area distinguish polymorphism of a gaptoglobin, ceruloplasmin and a group-specific component.

Gaptoglobin (Nr) has ability to enter actively connection with the hemoglobin dissolved in serum and to form the Hb complex — Nr. Consider that the molecule of the last owing to the big sizes does not pass through kidneys and, thus, gaptoglobin keeps hemoglobin in an organism. His main is seen in it fiziol, function (see. Gaptoglobin ). Assume that the enzyme of a gemalfametiloksigenaz splitting a protoporphyrinic ring on a α-methylene bridge to the bridge affects generally not hemoglobin, and on a complex of Hb-HP, i.e. usual exchange of hemoglobin includes its connection with Hp.

Determination of content of a gaptoglobin in blood serum matters for early diagnosis of some hron, diseases, for clarification of the reason of anemias, for clarification of the forecast and for establishment of efficiency of their treatment.

Fig. 1. Groups of a gaptoglobin (Nr) and the elektroforegramma characterizing them: each of groups of a gaptoglobin has the specific elektroforegramma differing in an arrangement, intensity and quantity of strips; on the right the corresponding groups of a gaptoglobin are designated; the minus sign designated the cathode, by a plus — the anode; the arrow at the word «start» designates an injection site of the studied serum in starched gel (for definition of its group of a gaptoglobin).
Fig. 2. Schemes immunoelektroforegramm groups of system G with: each of groups of the has system a specific arrangement and the form of an arch of precipitation with the corresponding antigens (are designated by figures) on Immunoelektroforezamm; 1 — albumine; 2 — a α-glycoprotein;;;;;;;;;; 3 — Gc 1; 4 — α22М; 5 — Gc 2; 6 — transferrin; the plus designated the anode; a minus sign — the cathode; circles — sites of application of serums.
Fig. 3. Schemes immunoelektroforegramm groups of transferrin at their research in starched gel: each of groups of transferrin (black strips) is characterized by various arrangement on an immunoelektroforegramma; letters over (under) strips designated various groups of transferrin (Tf); shaped strips correspond to an arrangement of albumine and a gaptoglobin (Hp).

In 1955 Smitis (O. of Smithies) established three basic groups of gaptoglobin which depending on electrophoretic mobility designate Hp 1-1, Hp 2-1 and Hp of 2-2 (fig. 1). Except these groups, other kinds of a gaptoglobin seldom meet: Nr2-1 (fashion), HpCa, Hp Johnson-type, Nr of Johnson Mod 1, Nr of Johnson Mod 2, type F, type D, etc. Seldom at people gaptoglobin is absent — an agaptoglobinemiya (Nr 0 — 0).

Groups of a gaptoglobin meet various frequency at persons of various races and nationalities. E.g., the HP group 2-1 — 49,5% most often occurs at the Russian population, the HP group 2-2 — 28,6% and the HP group 1-1 — 21,9% is more rare. The HP group 2-2 — 81,7% most often occurs at residents of India, on the contrary, and the HP group 1-1 makes only 1,8%. The population of Liberia has the HP group 1-1 more often — 53,3% and seldom the HP group 2-2 — 8,9%. The HP group 1-1 occurs at the population of Europe in 10 — 20% of cases, the HP group 2-1 — in 38 — 58%, and group of Nr 2-2 — in 28 — 45%.

Ceruloplasmin (Wednesday). It is described in 1961 by Owen and Smith (J. Owen, R. Smith). 4 groups differ: SRA, SRAV, SRV and SRVS. Most often the SRV group meets. At Europeans this group occurs in 99%, and at Negroids — in 94%. The SRA group at Negroids takes place in 5,3%, and at Europeans — in 0,006% of cases.

The group-specific component (Gc) is described in 1959. J. Hirschfeld. By means of an immunoelectrophoresis distinguish three basic groups — Gc 1-1, Gc 2-1 and Gc 2-2 (fig. 2). Very seldom other groups meet: Gc 1-X, Gcx-x, GcAb, Gcchi, Gc 1-Z, Gc 2-Z, etc.

The Gc groups meet unequal frequency at various people. So, among residents of Moscow the Gc 1-1 type makes 50,6%, Gc of 2-1 — 39,5%, Gc of 2-2 — 9,8%. There are populations among which the Gc 2-2 type does not occur. The Gc 1-1 type, and in 16,7% — the Gc 2-1 type, in 0,6% — the Gc 2-2 type occurs at residents of Nigeria in 82,7% of cases. Indians (Novayo) almost everything (95,92%) treat the Gc 1-1 type. At most of the European people frequency like Gc 1-1 fluctuates within 43,6 — 55,7%, Gc 2-1 — within 37,2 — 45,4%, Gc 2-2 — within 7,1 — 10,98%.

Globulins. Transferrin, post-transferrin and the 3rd component of a complement concern to them (β1c-глобулин). Many authors consider that post-transferrin and the third component of a complement of the person are identical.

Tf easily enters connection with iron. This connection easily breaks up. The specified property of transferrin provides performance to them important fiziol, functions — transfer of iron of plasma in the deionized form and its delivery in marrow where it is used at a hemopoiesis.

Transferrin has numerous groups: TfC, TfD, TfD1, TfD0, TfDchi, TfB0, TfB1, TfB2, etc. (fig. 3). Tf is available almost for all people. Other groups meet seldom and are distributed unevenly at various people.

Pt. Its polymorphism was described in 1969 by Rose and Gezerik (M. of Rose, G. Geserik). Distinguish the following groups of post-transferrins: And, AB, B, BC, C, EXPERT. At it. the population of group of post-transferrins meet the following frequency: And — 5,31%, AV — 31,41%, B-60,62 of %, BC-0,9 of %, With — 0%, AC-1,72 of %.

Third component of a complement (C '3). 7 groups C' 3 are described. They are designated or figures (C '3 1—2, C'3 1—4, C'3 1—3, C'3 1 —1, C'3 2—2, etc.), or letters (C' 3 S—S, C'3 F—S, C'3 F-F, etc.). At the same time 1 corresponds to the letter F, 2—S, 3 — So, 4 — S.

Lipoproteids. Distinguish three group systems designated by Ag, Lp and Ld.

In the Ag system Ag (a), Ag (x), Ag (b), Ag (y), Ag (z), Ag (t) and Ag(a1) antigens are found. B the Lp system enter Lp (a) and Lp (x) antigens. These antigens with various frequency occur at persons of different nationalities. Frequency of a factor of Ag (a) at Americans (white) — 54%, Polynesians — 100%, Micronesias — 95%, Vietnamese — 71%, Poles — 59,9%, Germans — 65%.

Various combinations of antigens also with an unequal frequency occur at persons of different nationalities. E.g., the Ag group (x — at +) at Swedes occurs in 64,2%, and at Japanese — in 7,5%, the Ag group (x+y —) for Swedes is available in 35,8%, and for Japanese — in 53,9%.

Blood groups in the medicolegal relation

the Research G. to. it is widely used in forensic medicine at the solution of questions of a doubtful paternity, motherhood (see. Motherhood disputable , doubtful paternity ), and also at a blood analysis on material evidences (see). Define group accessory of erythrocytes, group antigens of serumal systems and group properties of enzymes of blood.

Group accessory of blood of the child is compared with group accessory of blood of alleged parents. At the same time investigate the fresh blood received from these persons. The child can have only those group antigens which are available at least for one parent, and it belongs to any group system. E.g., mother has a blood group And, at the father — And, at the child — AV. The child with such G. to. could not be born from this couple since at this child one of parents shall have antigen B in blood.

For the same purposes antigens of the system MNSs, P, etc. E.g. are investigated, at a research of antigens of the R h system blood of the child does not may contain antigens Rho (D), rh' (C), rh" (E), hr' (e) and hr" (e) if this antigen is not in blood at least one of parents. The same treats antigens of the Duffy system (Fya — Fyb), the Kell system (To — k). Than more group systems of erythrocytes are investigated at the solution of questions of replacement of children, a doubtful paternity etc., that a high probability of receiving a positive take. Existence in blood of the child of the group antigen which is absent in blood of both parents at least on one group system is the undoubted sign allowing to exclude estimated paternity (or motherhood).

Also these issues at inclusion in examination of definition of group antigens of proteins of plasma are resolved — Gm, by Nr, Gc, etc.

In the solution of these questions begin to use definition of group signs of leukocytes, and also group differentiation of fermental systems of blood.

For the solution of a question of a possibility of an origin of blood on material evidences from the particular person also define group properties of erythrocytes, serumal systems and group distinctions of enzymes. At a research of spots of blood often determine antigens of the following izosero by l. systems: AB0, MN, P, Le, Rh. For G.'s definition to. in spots resort to special methods of a research.

Agglutinogens izosero l. systems can be found in spots of blood by use of the corresponding serums by various methods. In forensic medicine most often resort for these purposes to reaction of absorption in quantitative modification, absorption elution and the mixed agglutination.

The method of absorption is that previously define a caption of the serums entered into reaction. Then serums enter into contact with the material taken from a spot of blood. Through a nek-swarm time of serum is sucked away from a spot of blood and again titrated. On decrease in a caption of this or that applied serum judge existence in a spot of blood of the corresponding antigen. E.g., the spot of blood considerably lowered a caption of serum anti-In and Anti-R, therefore, in the studied blood there are antigens B and P.

Reactions of absorption elution and the mixed agglutination apply to identification of group antigens of blood especially when on material evidences there are traces of blood of the small sizes. Before statement of reaction from the studied spot take one or several threads of material, with to-rymi and work. At identification of antigens of a row izosero l. systems blood on threads is fixed methyl alcohol. For identification of antigens of some systems of fixing it is not required: it can lead to decrease in absorbing properties of antigen. Threads place in the corresponding serums. If in blood on a thread there is a group antigen corresponding to antibodies of serum, then these antibodies will be absorbed by this antigen. Then the antibodies which remained free delete by washing of material. In a phase of elution (process, the return absorption) threads place in a suspension of the erythrocytes corresponding to the applied serum. E.g., if in a phase of absorption serum was applied and, add erythrocytes of group A if serum anti-Lea, respectively, the erythrocytes containing Le (a) antigen etc. was applied. Then make thermal elution at t ° 56 °. At this temperature of an antibody leave to the environment since their communication with antigens of blood is broken. These antibodies at the room temperature cause agglutination of the added erythrocytes that is considered at microscopy. If in the studied material there are no antigens corresponding to the applied serums, then in a phase of absorption of an antibody are not absorbed and removed during the washing of material. In this case in a phase of elution free antibodies are not formed, and the added erythrocytes are not agglutinated. Thus it is possible to establish existence in blood of this or that group antigen.

Reaction of absorption elution can be executed in various modifications. E.g., elution can be made in fiziol, solution. The phase of elution can be carried out on slide plates or in test tubes.

The method of the mixed agglutination in initial phases is carried out, as well as a method of absorption elution. Distinction is only made by the last phase. Instead of a phase of elution at a method of the mixed agglutination of a thread place on a slide plate in a drop of a suspension of erythrocytes (erythrocytes shall have the antigen corresponding to the serum applied in a phase of absorption) and through a nek-swarm time observe drug microscopically. If in the studied object there is an antigen corresponding to the applied serum, then this antigen absorbs antibodies of serum, and in the last phase the added erythrocytes will «stick» to a thread in the form of nails or a beads since they will be held by free valencies of antibodies of the absorbed serum. If in the studied blood there is no antigen corresponding to the applied serum, then absorption will not happen, and all serum will be removed during the washing. In this case in the last phase the above described picture is not observed, and free distribution of erythrocytes in drug is noted. The method of the mixed agglutination is approved by hl. obr. concerning the AB0 system.

At a research of the AB0 system, except antigens, investigate also agglutinins by method of cover glass. On slide plates place the pieces which are cut out from the studied spot of blood and add a suspension of standard erythrocytes of blood groups to them And, B and 0. Drugs cover with cover glasses. If in a spot there are agglutinins, then they, being dissolved, cause agglutination of the corresponding erythrocytes. E.g., in the presence in a spot of agglutinin and agglutination of erythrocytes And etc. is observed.

For control the material taken from a material evidence out of the site soiled by blood is investigated in parallel.

At examination at first investigate blood of the persons who are connected to the case. Then their group characteristic is compared to the group characteristic of the blood which is available on a material evidence. If blood of any person differs according to the group characteristic from blood on material evidences, then in this case the expert can absolutely reject a possibility of an origin of blood on a material evidence from this person. At coincidence of the group characteristic of blood at any person and on material evidences the expert does not draw the categorical conclusion since he cannot reject in this case a possibility of an origin of blood on material evidences and from other person, blood to-rogo contains the same antigens.

See also Blood .

Bibliography: Boyd U. Fundamentals of immunology, the lane with English, M., 1969; Zotikov E. A., Manishkina R. P. and Kandelaki M. G. Antigen of new specificity in granulocytes, Dokl. Academy of Sciences of the USSR, it is gray. biol., t. 197, No. 4, page 948, 1971, bibliogr.; P. N jambs. Isoantigens and isoantibodies of the person it is normal also of pathology, M., 1974, bibliogr.; The application guide of blood and blood substitutes, under the editorship of A. N. Filatov, page 23, L., 1973, bibliogr.; Tumanov A. To, Bases of forensic medical examination of material evidences, M., 1975, bibliogr.; Tumanov A. K. and T about m silt and V. V N. Hereditary polymorphism of isoantigens and enzymes of blood is normal also of pathology of the person, M., 1969, bibliogr.; Umnova M. A. and Urinson R. M. About kinds of a Rhesus factor and their distribution among the population of Moscow, Vopr, antropol., century 4, page 71, 1960, bibliogr.; The unified methods of clinical laboratory trials, under the editorship of V. V. Menshikov, century 4, page 127, M. 1972, bibliogr.; Blood group immunology and transfusion techniques, ed. by J. W. Lockyer, Oxford, 1975; Blood and tissue antigens, ed. by D. Aminoff, p. 17, 187, 265, N. Y. — L., 1970, bibliogr.; Boorm a n K.E. Dodd B.E. An introduction to blood group serology, L., 1970; Fagerhol M. K.a. BraendM. Serum prealbumin, polymorphism in man, Science, v. 149, p. 986, 1965; Giblett E. R. Genetic markers in human blood, Oxford — Edinburgh, 1969, bibliogr.; Histocompatibility testing, ed. by E. S. Cur-toni a. o., p. 149, Copenhagen, 1967, bibliogr.; Histocompatibility testing, ed. by P. I. Terasaki, p. 53, 319, Copenhagen, 1970, bibliogr.; Klein H. Serumgruppe Pa/Gc (Postalbumin — group specific components), Dtsch. Z. ges. gerichtl. Med., Bd 54, S. 16, 1963/1964; Landstei-n e r K. t)ber Agglutinationserscheinungen normalen menschlichen Blutes, Wien. klin. Wschr., S. 1132, 1901; Landsteiner K. Levine P. A new agglutinable factor differentiating individual human bloods, Proc. Soc. exp. Biol. (N. Y.), v. 24, p. 600, 1927; Landsteiner K. Wiener A. S. Agglutinable factor in human blood recognized by immune sera for rhesus blood, ibid., v. 43, p. 223, 1940; M o r g a n W. T. J. Human blood-group specific substances, in book: Immunchemie, ed. by O. Westhphal, V. a. o., p. 73, 1965, bibliogr.; O w e n J. A. a. Smith H. Detection of ceruloplasmin after zone electrophoresis, Clin. chim. Acta, v. 6, p. 441, 1961; P a y n e R. o. A new leukocyte isoantigen system in man, Cold Spr. Harb. Symp. quant. Biol., v. 29, p. 285, 1964, bibliogr.; Procop O. u. Uhlen-b of of u with k G. Lehrbuch der menschlichen Blut-und Serumgruppen, Lpz., 1966, Bibliogr.; R a with e R. R. a. S a n g e r R. Blood groups in man, Oxford — Edinburgh, 1968; S h u 1 m a n N. R. a. o. Complement fixing isoantibodies against antigens common to platelets and leukocytes, Trans. Ass. Amer. Phycns, v. 75, p. 89, 1962; van der W e-erdt Ch. M. a. Lalezari P. Another Example of isoimmune neonatal neutropenia due to anti-Nal, Vox Sang., v. 22, p. 438, 1972, bibliogr.

P.N. Kosyakov; E. A. Zotikov (leukocytic groups), A. K. Tumanov (court. medical), M. A. Umnova (mt. issl.).