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Johnson HH. [[Paper - Twins, normal and abnormal|Twins, normal and abnormal]]. (1939) NY Acad. Sci. 1(4):
 
TRANSACTIONS
of THE NEW YORK ACADEMY OF SCIENCES
 
Ser. II, Vol. 1 FEBRUARY, 1939 No. 4
 
SECTION OF BIOLOGY‘
 
JANUARY 9, 1939


Doctor H. Herbert Johnson, College of the City of New York:
Doctor H. Herbert Johnson, College of the City of New York:


Twins, Normal and Abnormal.
(This lecture Jan 9, 1939 was illustrated by lantern slides, clay models, and specimens.)
 
(This lecture was illustrated by lantern slides, clay models, and specimens.)
 
In spite of the familiarity of the topic, it is obvious that
much is still to be learned concerning the exact origin in embryo of human twins and partial twins, while at the same time
there exist widespread misconceptions regarding the phenomenon of twinning. A review of current ideas bearing upon the
problem seems timely, especially since opportunity is given to
illustrate some points with preserved specimens of unusual
interest. No attempt is made here to present very original
points of view on the matter, and the discussion is limited to
well known theories of special interest to the embryologist.


Dizygotic (“fraternal” or two-egg) human twins theoretically may result from any one of three possibilities: 1. More or
less simultaneous ovulation from both ovaries; 2. Ovulation, at
about the same time, of more than one follicle in a single ovary;
3. Ovulation of a single follicle containing two or more ova.
Evidence is presented that such biovate or polyovate follicles
are much more frequent than is generally believed. The exhib
1 No meetings were held in January by the Sections of Geology and Anthropology.


TRANSACTIONS of the New York Academy of Sciences, Series II, Volume
In spite of the familiarity of the topic, it is obvious that much is still to be learned concerning the exact origin in embryo of human twins and partial twins, while at the same time there exist widespread misconceptions regarding the phenomenon of twinning. A review of current ideas bearing upon the problem seems timely, especially since opportunity is given to illustrate some points with preserved specimens of unusual interest. No attempt is made here to present very original points of view on the matter, and the discussion is limited to well known theories of special interest to the embryologist.
1, No. 4, February, 1939.


This publication is distributed to Members and is published monthly from
Dizygotic (“fraternal” or two-egg) human twins theoretically may result from any one of three possibilities: 1. More or less simultaneous ovulation from both ovaries; 2. Ovulation, at about the same time, of more than one follicle in a single ovary; 3. Ovulation of a single follicle containing two or more ova. Evidence is presented that such biovate or polyovate follicles are much more frequent than is generally believed. The exhib 1 No meetings were held in January by the Sections of Geology and Anthropology.
November to June, inclusive, at N. Queen St. and McGovern Ave., Lancaster,
Pa., by The New York Academy of Sciences, Seventy-ninth Street and Central
Park West, New York City.


Associate Editor: Eunice Thomas Miner, Executive Secretary.
This publication is distributed to Members and is published monthly from November to June, inclusive, at N. Queen St. and McGovern Ave., Lancaster, Pa., by The New York Academy of Sciences, Seventy-ninth Street and Central Park West, New York City.


Entered as second-class matter December 2, 1938, at the post ofice at Lancaster, Pa., under the act of August 24, 1912.


53
its include examples from the pig, rabbit, dog, cat and monkey. The view of Maximov that such follicles are characteristic of young females, is not supported by this investigation.
54 TRANSACTIONS


its include examples from the pig, rabbit, dog, cat and monkey.
It is now quite generally accepted that about one fourth of all twins are monozygotic (“identical” or single-egg) but no single line of evidence gives irrefutable proof that monozygotic twins exist. The various sources of evidence which collectively tend to establish proof of single-egg twins are reviewed in the recent book by Newman et al.
The view of Maximov that such follicles are characteristic of
young females, is not supported by this investigation.


It is now quite generally accepted that about one fourth of
Statistical evidence is gleaned from the fact that same-sexed twins occur more frequently than opposite-sexed twins, whereas the genetic expectation is a 1: 1 ratio of same-sexed to oppositesexed pairs. The weakness in this line of evidence when considered alone lies in the assumption that opposite-sexed twins are equally viable in utero as same-sexed pairs, and are in no way subject to environmental sex changing disturbances such as have been demonstrated experimentally in other higher vertebrates.
all twins are monozygotic (“identical” or single-egg) but no
single line of evidence gives irrefutable proof that monozygotic
twins exist. The various sources of evidence which collectively
tend to establish proof of single-egg twins are reviewed in the
recent book by Newman et al.


Statistical evidence is gleaned from the fact that same-sexed
Very important is the fact that conjoined twins are always of the same sex, and so are twins born in a single chorion. Until 1928 it was generally believed that all monozygotic twins develop in a common chorionic envelope. In 1927 Komai stated: “Apart from the examination of the placenta and foetal membranes at birth, there is no safe criterion of distinguishing the identical twins from the fraternal twins.””
twins occur more frequently than opposite-sexed twins, whereas
the genetic expectation is a 1: 1 ratio of same-sexed to oppositesexed pairs. The weakness in this line of evidence when considered alone lies in the assumption that opposite-sexed twins
are equally viable in utero as same-sexed pairs, and are in no
way subject to environmental sex changing disturbances such as
have been demonstrated experimentally in other higher vertebrates.


Very important is the fact that conjoined twins are always
The fact that identical twins are so frequently monochorial refutes a widespread dogma that such individuals originate by a splitting apart of the blastomeres at first cleavage. This point of View results from a misguided application to the human egg of the classical experimental results obtained on other forms. Chabry, Herbst, Driesch, Wilson et al. obtained more or less “normal” development of certain marine invertebrates and of Amphioxus from single blastomeres isolated experimentally during early cleavage. It should be emphasized that
of the same sex, and so are twins born in a single chorion. Until 1928 it was generally believed that all monozygotic twins
develop in a common chorionic envelope. In 1927 Komai
stated: “Apart from the examination of the placenta and foetal
membranes at birth, there is no safe criterion of distinguishing
the identical twins from the fraternal twins.””


The fact that identical twins are so frequently monochorial
1 Newman, Freeman, and Holzinger. “Twins: A Study of Heredity and Environment.” University of Chicago Press, 1937. 2Komai, Taku. Science, 65: 280. 1927.  
refutes a widespread dogma that such individuals originate by
a splitting apart of the blastomeres at first cleavage. This
point of View results from a misguided application to the human
egg of the classical experimental results obtained on other
forms. Chabry, Herbst, Driesch, Wilson et al. obtained more
or less “normal” development of certain marine invertebrates
and of Amphioxus from single blastomeres isolated experimentally during early cleavage. It should be emphasized that


1Newman, Freeman, and Holzinger. “Twins: A Study of Heredity and
these eggs do not self-isolate their own blastomeres. There may be no theoretical reason why the human ovum, or that of any mammal, might not divide at a stage in early cleavage into entirely separated blastomeres each capable of producing a complete body. The presence of the single chorionic envelope, however, shows that this is not usually the case. Since the chorion in the mammal embryo is developed from the outer layer of cells of the blastula and the embryo results from a clump of more centrally placed cells, it is obvious that the real division of the body in such cases results from the separation of this inner cell mass at a comparatively advanced stage of blastula formation. Such duplicated inner cell masses have been described in the armadillo by Patterson, the sheep by Assheton, and the.human by Arey.
Environment.” University of Chicago Press, 1937.
2Komai, Taku. Science, 65: 280. 1927.
THE NEW YORK ACADEMY or SCIENCES 55


these eggs do not self-isolate their own blastomeres. There
Since 1928 some evidence has accumulated to show that perhaps a few monozygotic twins are indeed dichorial. The case of identical triplets described by Komai and Fukuoko,‘ 1931, is not altogether conclusive when considered alone. Two bodies were produced in a chorion, and the third was in a separate chorion attached to the first. Komai interpreted this as a fusion of two separate chorions, but it could almost equally be called a faulty separation of one. Other evidence for dichorial identical twinning is presented by Curtius° and Lassen.’ It is still unknown whether these cases really represent a separation of early blastomeres, or a constriction of the chorion at a much later stage, perhaps while invasion of the uterine wall is taking place.
may be no theoretical reason why the human ovum, or that of
any mammal, might not divide at a stage in early cleavage into
entirely separated blastomeres each capable of producing a
complete body. The presence of the single chorionic envelope,
however, shows that this is not usually the case. Since the
chorion in the mammal embryo is developed from the outer
layer of cells of the blastula and the embryo results from a
clump of more centrally placed cells, it is obvious that the real
division of the body in such cases results from the separation
of this inner cell mass at a comparatively advanced stage of
blastula formation. Such duplicated inner cell masses have
been described in the armadillo by Patterson, the sheep by
Assheton, and the.human by Arey.


Since 1928 some evidence has accumulated to show that perhaps a few monozygotic twins are indeed dichorial. The case
Duplication of body parts occurs in an extensive series ranging from duplication of a single digit or the palm of the hand to duplication of the entire limb, the head, or practically the entire body (the conjoined, or “Siamese” twin). Conjoined twins apparently arise later in gastrulation than identical twins. The dissimilarity of the individuals has been explained in a view advanced by Newman.One is developed entirely from that part of the embryonic disc previously differentiated as left, and the other from the corresponding rudiment already determined as right. Each rudiment restores the missing portion in mirror-image symmetry to itself. Thus the two bodies differ to about the same degree as is frequently seen in the left and right sides of an ordinary person.
of identical triplets described by Komai and Fukuoko,1931, is
not altogether conclusive when considered alone. Two bodies
were produced in a chorion, and the third was in a separate
chorion attached to the first. Komai interpreted this as a
fusion of two separate chorions, but it could almost equally be
called a faulty separation of one. Other evidence for dichorial
identical twinning is presented by Curtius° and Lassen.’ It is
still unknown whether these cases really represent a separation
of early blastomeres, or a constriction of the chorion at a much
later stage, perhaps while invasion of the uterine wall is taking
place.


Duplication of body parts occurs in an extensive series ranging from duplication of a single digit or the palm of the hand
3 Arey, L. B. Anat. Rec., 23, 1922.  
to duplication of the entire limb, the head, or practically the
entire body (the conjoined, or “Siamese” twin). Conjoined


3Arey, L. B. Anat. Rec., 23, 1922.
4 Arey, L. B. “Developmental Anatomy,” 3rd Ed., p. 154, 1934. W. B. Saunders, Publisher.
4Arey, L. B. “Developmental Anatomy,” 3rd Ed., p. 154, 1934. W. B.
Saunders, Publisher.


5Komai, Taku and Gore Fukuoka. Jour. of Hered., 22, 1931.
5 Komai, Taku and Gore Fukuoka. Jour. of Hered., 22, 1931.


6 Curtius, F. Zeitschrift fiir Konstitutionslehre, 13, 1928.
6 Curtius, F. Zeitschrift fiir Konstitutionslehre, 13, 1928.


7 M. T. Lassen. Archiv fiir Gynakologie, 147, 1931.
7 M. T. Lassen. Archiv fiir Gynakologie, 147, 1931.
56 TRANSACTIONS


twins apparently arise later in gastrulation than identical
After reviewing various theories as to the basic causative factor leading to embryonic axial duplication, the theory of developmental arrest, advanced especially by Stockard,° still seems to be, in a general way, the best interpretation. The Spemann school has shown that the portion of the embryonic disc capable of being organized to produce a head is a considerably larger field than is actually necessary to produce a head. The center is the more responsive portion of this field. If a developmental arrest occurs precisely at the time that evocatory stimulation, capable of organizing a head, is expected, the center of the field is retarded and loses its power of development. If the retarding factor then ceases to operate, some power of organization still is retained by the more lateral portions of the head-field, and these separately begin development, thus forming a duplicated head. Similar arrest, occurring at the critical moment at which gastrulation should begin, might similarly result in the appearance of two gastrulation points on a single embryonic disc. The degree of mechanical and physiological separation of these centers is in direct ratio to the degree of separation exhibited by the resulting twins.
twins. The dissimilarity of the individuals has been explained
in a view advanced by Newman.‘ One is developed entirely
from that part of the embryonic disc previously differentiated
as left, and the other from the corresponding rudiment already
determined as right. Each rudiment restores the missing portion in mirror-image symmetry to itself. Thus the two bodies
differ to about the same degree as is frequently seen in the left
and right sides of an ordinary person.


After reviewing various theories as to the basic causative
factor leading to embryonic axial duplication, the theory of
developmental arrest, advanced especially by Stockard,° still
seems to be, in a general way, the best interpretation. The
Spemann school has shown that the portion of the embryonic
disc capable of being organized to produce a head is a considerably larger field than is actually necessary to produce a head.
The center is the more responsive portion of this field. If a
developmental arrest occurs precisely at the time that evocatory stimulation, capable of organizing a head, is expected, the
center of the field is retarded and loses its power of development. If the retarding factor then ceases to operate, some
power of organization still is retained by the more lateral portions of the head-field, and these separately begin development,
thus forming a duplicated head. Similar arrest, occurring at


the critical moment at which gastrulation should begin, might
Specimens exhibited included a foetal pig with duplicated snout and awe, and three eyes; a human foetus of five months with two complete heads, tour shoulders, and three arms; a human octopus. with two nearly complete bodies joined at the pelvis. These specimens and others shown. are part of the collection of the Division of Embryology of the College of the City of New York.


similarly result in the appearance of two gastrulation points
on a single embryonic disc. The degree of mechanical and
physiological separation of these centers is in direct ratio to
the degree of separation exhibited by the resulting twins.


Specimens exhibited included a foetal pig with duplicated snout and awe, and three
8 Newman, H. H. Journ. of He:-ed., 22, 1931. 9 Stockard, C. R. Amer. Jour. Anat., 28, 1921.
eyes; a human foetus oi.’ five months with two complete heads, tour shou ders, and three
arms; a human octopus. with two nearly complete bodies Joined at the pelvis. These
s cimens and others shown. are part 0 the collection of the Division of Embryology
o the College of the City of New York.


8 Newman, H. H. Journ. of He:-ed., 22, 1931.
9 Stockard, C. R. Amer. Jour. Anat., 28, 1921.


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Johnson HH. Twins, normal and abnormal. (1939) NY Acad. Sci. 1(4):

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This 1939 paper by Johnson describes early human embryo twinning.




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Twins, Normal and Abnormal

Doctor H. Herbert Johnson, College of the City of New York:

(This lecture Jan 9, 1939 was illustrated by lantern slides, clay models, and specimens.)


In spite of the familiarity of the topic, it is obvious that much is still to be learned concerning the exact origin in embryo of human twins and partial twins, while at the same time there exist widespread misconceptions regarding the phenomenon of twinning. A review of current ideas bearing upon the problem seems timely, especially since opportunity is given to illustrate some points with preserved specimens of unusual interest. No attempt is made here to present very original points of view on the matter, and the discussion is limited to well known theories of special interest to the embryologist.

Dizygotic (“fraternal” or two-egg) human twins theoretically may result from any one of three possibilities: 1. More or less simultaneous ovulation from both ovaries; 2. Ovulation, at about the same time, of more than one follicle in a single ovary; 3. Ovulation of a single follicle containing two or more ova. Evidence is presented that such biovate or polyovate follicles are much more frequent than is generally believed. The exhib 1 No meetings were held in January by the Sections of Geology and Anthropology.

This publication is distributed to Members and is published monthly from November to June, inclusive, at N. Queen St. and McGovern Ave., Lancaster, Pa., by The New York Academy of Sciences, Seventy-ninth Street and Central Park West, New York City.


its include examples from the pig, rabbit, dog, cat and monkey. The view of Maximov that such follicles are characteristic of young females, is not supported by this investigation.

It is now quite generally accepted that about one fourth of all twins are monozygotic (“identical” or single-egg) but no single line of evidence gives irrefutable proof that monozygotic twins exist. The various sources of evidence which collectively tend to establish proof of single-egg twins are reviewed in the recent book by Newman et al.‘

Statistical evidence is gleaned from the fact that same-sexed twins occur more frequently than opposite-sexed twins, whereas the genetic expectation is a 1: 1 ratio of same-sexed to oppositesexed pairs. The weakness in this line of evidence when considered alone lies in the assumption that opposite-sexed twins are equally viable in utero as same-sexed pairs, and are in no way subject to environmental sex changing disturbances such as have been demonstrated experimentally in other higher vertebrates.

Very important is the fact that conjoined twins are always of the same sex, and so are twins born in a single chorion. Until 1928 it was generally believed that all monozygotic twins develop in a common chorionic envelope. In 1927 Komai stated: “Apart from the examination of the placenta and foetal membranes at birth, there is no safe criterion of distinguishing the identical twins from the fraternal twins.””

The fact that identical twins are so frequently monochorial refutes a widespread dogma that such individuals originate by a splitting apart of the blastomeres at first cleavage. This point of View results from a misguided application to the human egg of the classical experimental results obtained on other forms. Chabry, Herbst, Driesch, Wilson et al. obtained more or less “normal” development of certain marine invertebrates and of Amphioxus from single blastomeres isolated experimentally during early cleavage. It should be emphasized that

1 Newman, Freeman, and Holzinger. “Twins: A Study of Heredity and Environment.” University of Chicago Press, 1937. 2Komai, Taku. Science, 65: 280. 1927.

these eggs do not self-isolate their own blastomeres. There may be no theoretical reason why the human ovum, or that of any mammal, might not divide at a stage in early cleavage into entirely separated blastomeres each capable of producing a complete body. The presence of the single chorionic envelope, however, shows that this is not usually the case. Since the chorion in the mammal embryo is developed from the outer layer of cells of the blastula and the embryo results from a clump of more centrally placed cells, it is obvious that the real division of the body in such cases results from the separation of this inner cell mass at a comparatively advanced stage of blastula formation. Such duplicated inner cell masses have been described in the armadillo by Patterson, the sheep by Assheton, and the.human by Arey.“

Since 1928 some evidence has accumulated to show that perhaps a few monozygotic twins are indeed dichorial. The case of identical triplets described by Komai and Fukuoko,‘ 1931, is not altogether conclusive when considered alone. Two bodies were produced in a chorion, and the third was in a separate chorion attached to the first. Komai interpreted this as a fusion of two separate chorions, but it could almost equally be called a faulty separation of one. Other evidence for dichorial identical twinning is presented by Curtius° and Lassen.’ It is still unknown whether these cases really represent a separation of early blastomeres, or a constriction of the chorion at a much later stage, perhaps while invasion of the uterine wall is taking place.

Duplication of body parts occurs in an extensive series ranging from duplication of a single digit or the palm of the hand to duplication of the entire limb, the head, or practically the entire body (the conjoined, or “Siamese” twin). Conjoined twins apparently arise later in gastrulation than identical twins. The dissimilarity of the individuals has been explained in a view advanced by Newman.‘ One is developed entirely from that part of the embryonic disc previously differentiated as left, and the other from the corresponding rudiment already determined as right. Each rudiment restores the missing portion in mirror-image symmetry to itself. Thus the two bodies differ to about the same degree as is frequently seen in the left and right sides of an ordinary person.

3 Arey, L. B. Anat. Rec., 23, 1922.

4 Arey, L. B. “Developmental Anatomy,” 3rd Ed., p. 154, 1934. W. B. Saunders, Publisher.

5 Komai, Taku and Gore Fukuoka. Jour. of Hered., 22, 1931.

6 Curtius, F. Zeitschrift fiir Konstitutionslehre, 13, 1928.

7 M. T. Lassen. Archiv fiir Gynakologie, 147, 1931.

After reviewing various theories as to the basic causative factor leading to embryonic axial duplication, the theory of developmental arrest, advanced especially by Stockard,° still seems to be, in a general way, the best interpretation. The Spemann school has shown that the portion of the embryonic disc capable of being organized to produce a head is a considerably larger field than is actually necessary to produce a head. The center is the more responsive portion of this field. If a developmental arrest occurs precisely at the time that evocatory stimulation, capable of organizing a head, is expected, the center of the field is retarded and loses its power of development. If the retarding factor then ceases to operate, some power of organization still is retained by the more lateral portions of the head-field, and these separately begin development, thus forming a duplicated head. Similar arrest, occurring at the critical moment at which gastrulation should begin, might similarly result in the appearance of two gastrulation points on a single embryonic disc. The degree of mechanical and physiological separation of these centers is in direct ratio to the degree of separation exhibited by the resulting twins.


Specimens exhibited included a foetal pig with duplicated snout and awe, and three eyes; a human foetus of five months with two complete heads, tour shoulders, and three arms; a human octopus. with two nearly complete bodies joined at the pelvis. These specimens and others shown. are part of the collection of the Division of Embryology of the College of the City of New York.


8 Newman, H. H. Journ. of He:-ed., 22, 1931. 9 Stockard, C. R. Amer. Jour. Anat., 28, 1921.



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