The 7 Sexes: Biology of Sex Determination

Views: 383
Ratings: (0)

Few of us know much about the biology of sex determination, but what could be more interesting than to discover how we are shaped into males and females? In this book, Elof Carlson tells the incredible story of the difficult quest to understand how the body forms girls and boys. Carlson's history takes us from antiquity to the present day to detail how each component of human reproduction and sexuality was identified and studied, how this knowledge enlarged our understanding of sex determination, and how it was employed to interpret such little understood aspects of human biology as the origin of intersex births.

List price: $29.99

Your Price: $23.99

You Save: 20%

 

22 Slices

Format Buy Remix

1 Introduction

ePub

With rare exceptions, animals consist of sexually reproducing populations that are roughly half male and half female—at least that is a human perspective that is applied to other mammals, and generalized to all other animals. An observant individual will notice roaches mating rear end to rear end or horseshoe crabs on the beach in springtime mating with the male mounted on a female, reinforcing the idea that the image of human intercourse can be generalized. I can observe fruit flies mating in the same way without use of a microscope, and I can even tell which is male and which is female if I am looking at a solitary fruit fly resting on my finger.

But that idea of universality is undermined if I observe copulating earthworms, which seem to be engaged in some sort of symmetrical mutual engagement. The ambiguity of the earthworm’s hermaphroditism is also present in most flowering plants. Students learn that pollen bearing stamens are present in the same flower with female components—assigned scholarly names like stigma, style, and ovary—but that is also not universal.

 

2 Wild Guesses in an Era of Scientific Ignorance

ePub

Almost all of the topics taught in K–12 or undergraduate introductory science courses come from work published in the last two centuries. Before the nineteenth century, very little of the chemistry, biology, geology, astronomy, or physics (other than Newtonian) that is covered in a twenty-first century class had been discovered. Almost all of a medical school curriculum, with the exception of gross anatomy, is a product of work done in the past two centuries. However, humanity centuries ago had the same curiosity about life and the universe as those born today. One of the universally recognized experiences of all people born is that roughly half are males and about half are females. When it comes to classifying who is a male and who is a female at birth, almost every adult in the world will use the external genitals. In a male there are a penis and a scrotum containing two testes. In a female there is a vaginal passageway surrounded by labia and a clitoris. Except for rare occasions we do not see the genitalia of our fellow adult human beings. Usually we classify a person as male or female by characteristics such as body shape, the presence or absence of hair on the face, the length, distribution, and style of hair from the cranial part of the head, the bony structure of the limbs and face, the deepness or higher pitch of voice, the presence or absence of enlarged breasts, and the presence or absence of an “Adam’s apple.” To that, in most cultures, we add the clothing that people wear and the behavior we assign to males or females out of tradition or experience.

 

3 The Ancient World

ePub

Cellular life, sex determination, reproduction, and the processes associated with gestation are largely unseen and unknowable without the tools of science. A major reason for this is the scale at which life takes place. Most of the fundamental processes occur at the level of cells, chromosomes, genes, and molecules, none of which are visible to the unaided eye. Even the concepts of cells, chromosomes, genes, and molecules were not known to the ancient world except in the vaguest theoretical or speculative ways (e.g., atoms as ultimate units that are indivisible, or atoms as vortices of energy or motion). In Greek philosophic writings, Democritus of Abdera (460–370 BCE) believed females arose from the left testis and males from the right testis. Pythagoras (580–500 BCE) believed the male semen gave rise to the “noble parts” of the body and the female semen gave rise to the “gross parts.” Hippocrates (460–377 BCE) referred to the “liquor” or “sperma” produced by a copulating couple, and he believed strong sperma from either sex would produce males, while weak sperma from either sex would produce females. The “liquors” were believed to represent the entire body of the individual at the time of copulation, a view not too far from Charles Darwin’s (1809–1882) idea of pangenesis in 1868.1

 

4 Monotheistic Religious Interpretations

ePub

Countries with substantial populations of Christians and Muslims follow a monotheistic tradition derived from the Jews and their scriptures, the Old Testament or Hebrew Bible. Sex determination of humans is introduced in the book of Genesis as two separate events: Adam is first created as a male and later given a female companion, Eve, derived from his rib. The first strange feature of this separate creation of the two human sexes is that it occurs after the creation of sea, air, and land animals. These creatures are created in an unspecified number and commanded to multiply their kind. By implication, male and female representatives of these other forms of life were created without comment about their having two sexes, as were species that produced their progeny by parthenogenesis, budding (or other cloning mechanism), multiple mating types (e.g., paramecia), or hermaphroditic mutual gametic exchange (e.g., earthworms and snails).

The second unusual feature of the Genesis account is the creation of a female, Eve, presumably having a 46,XX chromosomal composition from the rib of Adam, presumably having a 46,XY chromosomal composition. Barring some miraculous act and assuming the Creator was using an XY mechanism for sex determination, the rib would have had to contain this 46,XX tissue. This would mean that Adam was some sort of chromosomal mosaic (or chimera, if the XX tissue was some type of embedded twin), with a region of his body (the rib area used for Eve) containing a karyotype that required two separate, nondisjunctional events to bring about. If one attributes miraculous acts for this formation of Eve, why was the rib necessary? If the human female has her origin as a second sex from Adam, the chromosome difference has to be reconciled (at least for those calling themselves “Creationist scientists”). One such possibility would be the lagging of the Y shifting a cell to 45,X, followed by a delayed separation of chromatids of the X producing the 46,XX cell from which the rib area was derived. The two X chromosomes would necessarily be identical in nucleotide sequence (except for a few new spontaneous mutations).1 Note the coincidence (or possible association) between the hermaphroditic nature of Adam before Eve is extracted from his rib, and the Greek myth cited by Plato in The Symposium, where heterosexual couples were produced from an initially hermaphroditic state.

 

5 The Descriptive Embryology of Male and Female Development

ePub

The study of embryology was extremely limited until the 1860s, when several techniques came together. One was the development of achromatic lenses, which allowed scientists to observe cells or other small objects without the confusion of chromatic aberration and spherical aberration. Chromatic aberration involves the breaking up of white light into a rainbow of concentric colors around an object. Spherical aberration is even worse because there is no central focus and competing points of convergence lead to a blurred image. Joseph Jackson Lister (1786–1869), whose more famous son, Joseph Lister (1827–1912), introduced antiseptic surgery, provided the solution.1

Like his father, Lister was a wine merchant. They were Quakers and fairly pious. Lister worked after hours on his interest in natural history. When he took up microscopy, he found the interference with sharp images annoying and switched to making his own lenses. Lister found that dissolving lead and other salts into glass changed the density within these glasses, which yielded different focal lengths when he made lenses from them. He eventually worked out a combination of two layers of molten glass (crown glass and flint glass), one correcting the other for both chromatic and spherical aberration. The resulting achromatic lenses revolutionized both astronomy (allowing bigger telescopic lenses) and biology (allowing compound microscopes that would multiply an object up to 2000 times. He began his experiments on lenses in 1824 and found a suitable achromatic lens in 1826, which he refined until publishing his findings in 1830.

 

6 The Discovery of the Egg in Higher Eukaryotes

ePub

While it was known to almost all of humanity since antiquity that birds lay eggs, it was not known that the egg is a cell until cell theory was developed in the 1830s. In 1652, William Harvey (1578–1657) found two things of interest when studying reproduction in the fallow deer.1 One was that after copulation and conception in the fall, the gestation process of the fallow deer goes into an arrested state for several months. The process is called diapause and it allows seasonal regulation of when birth takes place. It occurs during the blastocyst stage and was observable to Harvey as a small spot that neither went away nor enlarged until the winter. Harvey’s other, more significant, observation was that after copulation the uterus of the deer was empty. There was no coagulum to be seen or evidence that anything had changed in the uterus. What Harvey did not know was he could not see the blastocyst that implants itself in the uterus of the fallow deer because it is transparent and microscopic. The mammalian blastocyst at implantation at best would be the size of a period in a sentence. It does not enlarge before implantation in the uterus. It receives no external nutrient to grow until it implants. In 1843, Theodor L. W. Bischoff (1807–1882) was able to demonstrate the presence of that blastocyst. Bischoff studied the egg in rabbits, dogs, guinea pigs, and the roe deer, following it through its cleavages, blastocyst formation, and early embryonic development.2

 

7 The Discovery of Sperm in Higher Eukaryotes

ePub

Semen has long been recognized as necessary for producing offspring. It is liquid, somewhat viscous, and usually clear or slightly cloudy in appearance; certainly the unaided eye can see no visible body within it. The Greeks, especially through Hippocrates and later Galen, embraced a theory of vital fluids, which they called humors. Blood was considered the major constituent of life, at least among vertebrates. It was considered the progenitor of semen in the male body, and believed to be the hereditary material that allowed a species to generate offspring in its likeness.

Semen was endowed with a capacity to impose form on the pliable material supplied by females. That material was also thought to be blood: sometimes it was associated with menstrual blood, and sometimes it was thought to be another type of semen. Female semen was not clarified, like male semen, but still bloodlike and clotted—a type of miniscule clay ready to be molded into shape by the empowering effect of male semen. For more than two thousand years, arguments were made about the relative roles that males and females play in forming a new individual through their fluids, which were commingled after copulation. There were inside–outside theories in which the male supplied the outer components of the new baby. There were theories in which the female role was passive, being shaped exclusively by the male, forcing some observable phenomena, such as the equal contributions made to the skin color of the offspring of a black person and a white person, to be swept under a mental rug.

 

8 The Discovery of Sex Hormones

ePub

In 1902, William Bayliss (1860–1924) and Ernest Starling (1866–1927) introduced the term “hormone.”1 Hormones are substances produced by one organ, an endocrine gland, that acts at a distance on another organ. The field of science that studies this is called endocrinology. The names of hormones were all coined in the twentieth century, but the idea that there was something like hormones has existed since antiquity. For example, Chinese medicine frequently made use of extracts from human urine that were used to treat disease.

Since human history began and medical treatments were attempted, physical changes associated with endocrine glands have been known. Castrated males, since antiquity known as eunuchs, lose their capacity to grow a beard, may develop enlarged breasts, and become effeminate. Eunuchs have had a long history serving as guards of harems in the Middle East, where plural marriages were common and reflective of wealth and power, and they served as political advisors in the Forbidden City in Beijing during the rule of Chinese emperors. Eunuchs were usually castrated as young men, but a special category of eunuchs were castrated as preadolescent boys. These were called castrati. During the Renaissance and until the eighteenth century, boys in choirs who were aged six to ten and who had a talent for singing and reading music were castrated and groomed to become prized singers because of their “celestial” soprano-like upper voice range. They differed from typical eunuchs, who lost their testes as adults, tending to be taller than average and appearing “etiolated,” with unusually wide hips in an otherwise slender frame.2 Castration was also applied to slaves in Greece about 400 BCE because they were considered to be more docile. In Jewish tradition, eunuchs were excluded from religious ceremonies. Early Christian monks sometimes practiced castration to remove the temptation of sexual attraction.3

 

9 Ploidy Levels and Sex Determination

ePub

We tend to think of all animals as being diploid, represented as 2N, and their gametes as haploid, represented as N. Exceptions are rare: this is not true for only about 18 of the many thousands of taxonomic groups of animals. In plants like mosses and liverworts, an alternation of haploid and diploid states is common and among a category of insects called the Hymenoptera, which includes bees, ants, and wasps the presence of both haploid and diploid organisms is virtually the rule. These insects share a common mode of sex determination and also a social structure, called eusociality, in which there is usually one egg-laying queen, a huge number of sterile female workers or helpers, and a small number of drones that inseminate the queen (in some species just a single male for just a single encounter serves that role). Bees have immense commercial value, fertilizing one third of the plants we consume and having been used for honey production since antiquity. They cannot be domesticated, however, and experiments to mate specific males with specific females failed until the 1940s, when people started using artificial insemination to initiate bee genetics.

 

10 The Discovery of Sex Chromosomes

ePub

The idea of the continuity of chromosomes arose among German cytologists in the 1880s. At that time, many American biologists would get their PhDs (especially at Johns Hopkins University) and take a trip to Europe to visit the laboratories of German, Belgian, Dutch, French or Italian biologists where much of the work on meiosis, mitosis, and reproductive biology was taking place. They would then return to the United States to begin their own cytological studies. In 1891, the German biologist Herman Henking (1858–1942) studied the fire wasp, Pyrrhochoris,1 which is not actually a wasp, or Hymenopteran, but a true bug, or Hemipteran. He noted that, during the spermatogenesis of the fire wasp, there is an unusual chromosome: a nucleolar object that takes on a very dark stain in the first meiotic division. In the second division, this unit did not divide, and it appeared to remain in only one of the two cells produced. Because it was unusual in its staining, its morphology, and its behavior, Henking called it an X element, using X as a mathematical symbol for an unknown to be solved. The next year, when Henking was given an opportunity to take on an important and more financially rewarding position in German fisheries, he dropped cytology, focusing on fisheries for the rest of his career. Henking made no association between his X element and sex determination.

 

11 The Balance Theory of Sex Determination

ePub

From 1907, when Thomas Hunt Morgan began working on fruit flies, until 1915, he and his students believed that their sex chromosome composition was 2N = 8,XX for females and 2N = 7,XO for males because they had misinterpreted a paper that Nettie Stevens wrote in 1907. Once Morgan and his students realized that Drosophila melanogaster used the XX female and XY male system for sex determination, they had to reconcile the role of the Y chromosome in sex determination.1 Since it was well known from Edmund B. Wilson’s and Stevens’s work that some Diptera had XO males and others had XY males, they concluded that the Y could not be playing a role in the sex determination of males.

That inference was reinforced when Calvin Blackman Bridges (1889–1938) discovered a phenomenon he called nondisjunction, the topic on which he wrote his PhD dissertation, which was published in 1916.2 Bridges found an unexpected appearance of a white-eyed male in a cross that should have given red-eyed males. If one parent is a white-eyed male and the female parent is red eyed, all the progeny should be red eyed. When Bridges tried to mate the white-eyed male, he found it was sterile. He also found that if he did a cross with a white-eyed female and red-eyed male, the offspring should be white-eyed sons and red-eyed daughters: a distribution that the laboratory referred to as crisscross inheritance. But Bridges found a female that was white eyed on some occasions. That exceptional female was fertile, and when mated to a red-eyed male she gave an unusual distribution of progeny: about eight percent of the offspring being of an unexpected kind with respect to their eye color and sex.

 

12 The Discovery of Sex in Microorganisms

ePub

When Anthony van Leeuenhoek observed the animalcules, as he called them, from different dips of water or from his own body, he did not discuss how they formed. Most of his contemporaries would have said that they formed from spontaneous generation. The idea is as old as written thought. Aristotle believed in spontaneous generation, and so did anyone watching rotting food or meat swarming with maggots. Before Rudolph Virchow and Robert Remak’s cell doctrine, biologists did not think of life coming from preexisting life. At least they conceived the process as far back as life goes: Genesis for the pious; after Charles Darwin, some sort of event that led to the formation of the first living cell; or after H. J. Muller, the formation of the gene, the first replicating molecule that could copy its errors.

Microscopy flourished in the last half of the nineteenth century. It spun off the field of histology in medical schools and the field of cytology that led to inquiries about heredity. It was a necessary tool for the field of microbiology that flowed from germ theory. Louis Pasteur (1822–1895) and Robert Koch (1843–1910) introduced the germ theory of infectious diseases in the 1870s and 1880s. It revealed even smaller organisms than those seen by Robert Hooke and Leeuenhoek. Pasteur and Koch’s theory brought microscopy back to the medical school to study infectious diseases caused by bacteria and other microorganisms. By the end of the nineteenth century, scientists inferred the existence of even smaller organisms, which slipped through filters that barred passage of bacteria. In 1892, the first virus, tobacco mosaic virus, was identified.1

 

13 The History and Interpretations of Hermaphrodites and Intersexes

ePub

The concepts of hermaphroditism, sexual chimerism, sexual mosaicism, intersexuality, and gynandromorphism are all connected through a common aspect. They involve the presence, in an individual, of the sexual phenotype of both sexes to some degree, either at once or at some time in their life cycle as adults. In the broadest sense, the term intersex is most inclusive, and it is purged of connotations of the classic mythology of Hermaphroditus and his encounter with Salmacis. Unfortunately, the biological use of the term intersex has usually excluded errors of mitosis leading to mosaicism. Richard Goldschmidt (1878–1958) first introduced the term “intersexuality” to genetics in 1915, when he was working on gypsy moth sex determination. Thomas Hunt Morgan and his students first used the term “gynandromorph” in genetics in 1914 to describe mosaicism, usually an XX/X somatic composition of flies that had lost an X-chromosome due to non-disjunction after fertilization.1

Intersexuality implies a genetic or environmental cause for the dual-sexual aspect, in which all the cells of the body have the same genotype and chromosome composition. The term “chimerism” is applied to a special type of mosaicism. It has a classical legend behind it: numerous deities were part human and part animal in Egyptian and Greek mythologies. The Egyptian Sphinx is a part-human, part-lion, chimera that is familiar to most of humanity. In human biology, the term chimerism refers to individuals produced from four gametes (“tetragametic”) with two separate fertilizations, a fusion of what should have been non-identical twins into a single individual. In contrast, biologists use the term mosaicism to describe individuals who have a mitotic, nondisjunctional event (or gene mutation after zygote formation), leading to two different genetic compositions or cell lines that were derived from one initial fertilization. In medicine and psychology, the term “disorder or difference of sexual differentiation” (DSD) is replacing all of these terms when discussing patients with quite varied modes of origin. Time will tell whether this term is specific enough or accurate enough to convey information about the condition, as this new usage is put into public discourse. It is unlikely that the term DSD will be applied to fruit flies or species other than humans. Such fine distinctions among these terms are almost entirely of twentieth-century origin, because neither the sex chromosomes nor genes in relation to sex determination were known before then.

 

14 Dosage Compensation and the Sex Chromosomes

ePub

Calvin Bridges and Thomas Hunt Morgan discovered the existence of dosage differences on the X chromosome of fruit flies when comparing the allele of white eyes called eosin to that of the allele of white eyes called apricot. They called the phenomenon bicolorism, but did not make a generalization about it. In a stock of eosin flies, females had a darker eye color than males. In a stock of apricot flies, the eye color of the male and the female was the same. Eosin arose in a bottle of white-eyed flies as a solitary male fly. It was interpreted as a partial reverse mutation from white to eosin. About a decade later, in 1926, Curt Stern (1902–1981) discovered a mutation called bobbed bristles. It was the first genetic character found on the Y chromosome in fruit flies that was not associated with fertility.1 As it turns out, the shorter and slightly elevated bristles are associated with a gene on both the X and the Y chromosome. This made the gene behave like an autosomal recessive. Normal males and females had two doses, but XO males had a single dose and XXY fertile females had a triple dose. Stern noted that as the number of bobbed alleles increases, there is a normalizing effect.

 

15 The Discovery of Human Sex Chromosome Conditions

ePub

When Michael Frederic Guyer (1874–1959) looked at human chromosomes in 1910, he estimated the diploid number was 24 in females and 23 in males.1 He thought the sex chromosome mechanism was XX female and XO male. Two years later, Hans Jean Chrysostome von Winiwater (1875–1949) doubled that estimate and claimed the human chromosome number was 47 in spermatocytes, but he agreed with Guyer about the XO status of males.2 That changed in 1921, when Theophilus Shickel Painter (1889–1969) at the University of Texas reported a chromosome number of 48 (although he said it could be 46 or 47). Painter used testes from freshly executed prisoners or from castrated patients in mental asylums. He also said there was a definite Y chromosome in the male cells he studied and thus 48,XY was the chromosome number and sexual status of males for another generation.3

The status remained stable because techniques did not change in cytology for human cells until the 1950s, when tissue culture techniques improved, hypotonic solutions were used to increase the volume of nuclei, and colchicine was used to arrest cell division at metaphase. In 1956, when two Swedish investigators, Joe Hin Tjio and Albert Levan, combined all these procedures, they got a consistent reading of 46 chromosomes, with very clear XX females and XY males.4 From then on, the human chromosome number was represented as 46,XX for females and 46,XY for males. By 1960, a standardized way of photographing, enlarging, and clipping out chromosomes was introduced. The chromosomes were measured and aligned in size, place, and by grouping within a particular size range of the location of the centromere that separates the two arms of the chromosomes. (The shorter arm of the chromosome is called “p” and the longer arm “q.”) The resulting mounted representation of the chromosomes is called a karyotype. The X chromosome is in the C group (chromosomes of moderate size), and it is slightly submetacentric. The Y chromosome is very small, and the q arm is at least twice as long as the p arm. In 1964, Lionel Sharples Penrose (1898–1972) estimated that the X accounted for about 6 percent of the total DNA of the sperm nucleus in humans, and the Y accounted for about 2 percent.5

 

16 The Seven Sexes of Humans

ePub

In 1958, when I was a freshly minted PhD from Muller’s laboratory at Indiana University, I took my first academic job at Queen’s University in Kingston, Ontario. There, I was asked to teach a course in human genetics for medical students, which was a challenge because I had never had a human genetics course. As an offshoot from my dissertation study on the structure of the dumpy locus in fruit flies, I had published an article on the parallel of that gene complex to the Rh blood groups. That was my only contact with human genetics. I spent a lot of time in the library at Indiana University, and at Queen’s University when I arrived there, reading what I could about human genetics, including Curt Stern’s pioneering text in this field.1 I hit the medical books and journals, looking for human parallels to genetic processes in fruit flies and other organisms. When I came to the topic of sex determination, I knew that fruit flies and humans both had XX female and XY male sex chromosomes. I read Jones and Scott’s fine text on hermaphroditic and pseudohermaphroditic disorders, and dipped into some human embryology texts to follow what was then known about sex differentiation in humans, both in normal and in clinically abnormal sexual development.2 I organized the information in my mind and presented it in a series of lectures that I called, at the time, “the seven sexes of man.”

 

17 The Identification and Role of Sex-Determining Genes

ePub

Life can be resilient and vulnerable at the same time. We rejoice at stories of Olympic medalists who overcome severe injury or a childhood marked with tragedy. At the same time, nature can dish out genetic disorders resulting in births incompatible with life—severely limiting in organ function or leaving an adult with chronic illness. What makes life so vulnerable is the nature of genetic material. Something as simple as altering or removing one nucleotide pair out of some three billion present in a sperm or an egg can result in one of those debilitating or lethal genetic conditions. That doesn’t happen if one pulls a single brick out of a multistoried building. It won’t collapse no matter where that brick is removed. If the gene happens to involve the sex-determining programs in the embryo, the results can be quite dramatic.

The sex-determining genes can be found on the sex chromosomes and the autosomes. This is no surprise, because many organ systems are involved in sex pathologies. They can result from abnormalities of the pituitary gland, the hypothalamus, the adrenal cortex, the gonads, or those embryonic structures that will form the internal and external genitalia. A major gene involved in male sex determination is the SRY gene. It is on the Y chromosome and located at Yp11.3, i.e., on the short or p arm of the Y chromosome at band region 11.3 (Figure 17.1). Andrew H. Sinclair found the SRY gene while he was working with Peter Goodfellow’s laboratory at Cambridge University in 1990.1 Prior to that, in 1987, David C. Page believed that what was called the “testes determining factor” (TDF) was a zinc finger gene in a different region.2 The TDF was known to be Y-associated from cytological studies of certain partial or complete sex reversals. Thus, a baby with X isoYp sex chromosomes has a duplication of the p arm. Such an individual is a male and sterile, because the q arm of the Y provides the genes associated with spermatogenesis, but otherwise functional. But a baby with X isoYq has a duplication of the q of the Y chromosome and an intact X chromosome and is missing the short arm of the Y chromosome. Because such babies are born as females with no testes, that is where the testes determining factor has to be located. The gonads of such babies are streaks, like those of babies with Turner syndrome. They also show other symptoms similar to those associated with Turner syndrome, suggesting that some growth factor is associated with the Yp region.

 

18 The History of Homosexuality

ePub

All human behavior is subject to the judgment of others. As children, we are judged by our parents, teachers, and playmates. Some behaviors are approved and admired; others are condemned and sometimes punished. That has been the history of behavior concerning table manners, dressing, grooming, reliability, dishonesty, theft, selfishness, generosity, cursing, bullying, flirting, and just about anything we do. Both culture and religion have their dos and don’ts. Those values change from generation to generation, and they are different in different countries and regions of countries. Almost all cultures condemn violent behavior toward those who are not designated by the state as legitimate objects for attack. We punish perjury, theft, fraud, treason, blackmail, piracy, and many other behaviors as crimes, and regulate them with laws. Most industrial nations no longer regard some crimes of the past as crimes today. At one time, blasphemy was a capital crime. Until the 1920s, it was a crime for a physician in the United States to offer medical advice on birth control. Until the 1950s, it was a crime for a white person to marry or live with a black person. Until the 1970s, a physician who carried out an abortion committed a crime.

 

Load more


Details

Print Book
E-Books
Slices

Format name
ePub (DRM)
Encrypted
true
Sku
2370004741451
Isbn
9780253006547
File size
0 Bytes
Printing
20 times / 30 days
Copying
20 times / 30 days
Read aloud
No
Format name
ePub
Encrypted
No
Printing
Allowed
Copying
Allowed
Read aloud
Allowed
Sku
In metadata
Isbn
In metadata
File size
In metadata