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GARDNER AND SUTHERLAND’S
CHROMOSOME ABNORMALITIES
AND GENETIC COUNSELING
2
OXFORD MONOGRAPHS ON MEDICAL GENETICS
General Editors:
JUDITH G. HALL PETER S. HARPER LOUANNE HUDGKINS EVAN EICHLER CHARLES J.
EPSTEIN (DECEASED 2011) ARNO G. MOTULSKY (RESIGNED 2011)
3
33. V. P. Sybert: Genetic skin disorders
34. M. Baraitser: Genetics of neurological disorders, third edition
35. H. Ostrer: Non- Mendelian genetics in humans
36. E. Traboulsi: Genetic factors in human disease
37. G. L. Semenza: Transcription factors and human disease
38. L. Pinsky, R. P. Erickson, and R. N. Schimke: Genetic disorders of human
sexual development
39. R. E. Stevenson, C. E. Schwartz, and R. J. Schroer: X- linked mental
retardation
40. M. J. Khoury, W. Burke, and E. Thomson: Genetics and public health in the
21st century
41. J. Weil: Psychosocial genetic counseling
42. R. J. Gorlin, M. M. Cohen, Jr., and R. C. M. Hennekam: Syndromes of the head
and neck, fourth edition
43. M. M. Cohen, Jr., G. Neri, and R. Weksberg: Overgrowth syndromes
44. R. A. King, J. I. Rotter, and A. G. Motulsky: Genetic basis of common
diseases, second edition
45. G. P. Bates, P. S. Harper, and L. Jones: Huntington’s disease, third edition
46. R. J. M. Gardner and G. R. Sutherland: Chromosome abnormalities and genetic
counseling, third edition
47. I. J. Holt: Genetics of mitochondrial disease
48. F. Flinter, E. Maher, and A. Saggar- Malik: Genetics of renal disease
49. C. J. Epstein, R. P. Erickson, and A. Wynshaw-Boris: Inborn errors of
development: The molecular basis of clinical disorders of morphogenesis
50. H. V. Toriello, W. Reardon, and R. J. Gorlin: Hereditary hearing loss and its
syndromes, second edition
51. P. S. Harper: Landmarks in medical genetics
52. R. E. Stevenson and J. G. Hall: Human malformations and related anomalies,
second edition
53. D. Kumar and S. D. Weatherall: Genomics and clinical medicine
54. C. J. Epstein, R. P. Erickson, and A. Wynshaw-Boris: Inborn errors of
development: The molecular basis of clinical disorders of morphogenesis,
second edition
55. W. Weber: Pharmacogenetics, second edition
56. P. L. Beales, I. S. Farooqi, and S. O’Rahilly: The genetics of obesity syndromes
57. P. S. Harper: A short history of medical genetics
58. R. C. M. Hennekam, I. D. Krantz, and J. E. Allanson: Gorlin’s syndromes of
the head and neck, fifth edition
59. D. Kumar and P. Elliot: Principles and practices of cardiovascular genetics
60. V. P. Sybert: Genetic skin disorders, second edition
61. R. J. M. Gardner, G. R. Sutherland, and L. C. Shaffer: Chromosome
abnormalities and genetic counseling, fourth edition
62. D. Kumar: Genomics and health in the developing world
63. G. Bates, S. Tabrizi, and L. Jones: Huntington’s disease, fourth edition
4
64. B. Lee and F. Scaglia: Inborn errors of metabolism: From neonatal screening
to metabolic pathways
65. D. Kumar and C. Eng: Genomic medicine, second edition
66. R. Stevenson, J. Hall, D. Everman, and B. Solomon: Human malformations
and related anomalies, third edition
67. R. Erickson and A. Wynshaw-Boris: Epstein’s inborn errors of development:
The molecular basis of clinical disorders of morphogenesis, third edition
68. C. Hollak and R. Lachmann: Inherited metabolic disease in adults: A clinical
guide
69. V. P. Sybert: Genetic skin disorders, third edition
70. R. J. M. Gardner and D. J. Amor: Gardner and Sutherland’s chromosome
abnormalities and genetic counseling, fifth edition
5
GARDNER AND SUTHERLAND’S
R. J. McKinlay GARDNER
ADJUNCT PROFESSOR
CLINICAL GENETICS GROUP
UNIVERSITY OF OTAGO, DUNEDIN, NEW ZEALAND
David J. AMOR
LORENZO AND PAMELA GALLI CHAIR
UNIVERSITY OF MELBOURNE
VICTORIAN CLINICAL GENETICS SERVICES
MURDOCH CHILDREN’S RESEARCH INSTITUTE
ROYAL CHILDREN’S HOSPITAL, MELBOURNE, AUSTRALIA
6
Oxford University Press is a department of the University of Oxford. It furthers the University’s
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All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or
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You must not circulate this work in any other form and you must impose this same condition on any
acquirer.
This material is not intended to be, and should not be considered, a substitute for medical or other
professional advice. Treatment for the conditions described in this material is highly dependent on
the individual circumstances. And, while this material is designed to offer accurate information with
respect to the subject matter covered and to be current as of the time it was written, research and
knowledge about medical and health issues is constantly evolving and dose schedules for
medications are being revised continually, with new side effects recognized and accounted for
regularly. Readers must therefore always check the product information and clinical procedures
with the most up-to-date published product information and data sheets provided by the
manufacturers and the most recent codes of conduct and safety regulation. The publisher and the
authors make no representations or warranties to readers, express or implied, as to the accuracy or
completeness of this material. Without limiting the foregoing, the publisher and the authors make
no representations or warranties as to the accuracy or efficacy of the drug dosages mentioned in the
material. The authors and the publisher do not accept, and expressly disclaim, any responsibility for
any liability, loss or risk that may be claimed or incurred as a consequence of the use and/or
application of any of the contents of this material.
7
This book is dedicated to Jocelyn, Geoffrey, and Craig, their
parents, and all other families who seek our “chromosomal
advice.”
8
for confirmatory advice about his low genetic risk.
9
Heredity
Inescapably, this is me—the diagnosis
is cause for anger at those
who brightly say we choose our destinies.
There is no store
of courage, wit or will
can save me from myself and I must face
my children, feeling like
that wicked fairy, uninvited
at the christening, bestowing on my own,
amidst murmurs of apprehension, a most
unwanted gift—that
of a blighted mind. No one
could tell me of this curse when I
was young and dreamt of children
and the graces they would bear. Later,
it seemed that a chill morning
revealed deeper layers
of truth. For my romancing
there is a price to pay—
perhaps my children’s children
will pass this tollgate after me.
My grandmothers gaze down from their frames
on my wall, sadly wondering.
—Meg Campbell
Dear DNA
In real life you’re just
a tangle of white filaments
captured in a test-tube,
and your first photo is not flattering:
grey smudges like tractor tracks,
or a rusty screw. Yet
many say you are beautiful.
Online for a night
with a hundred fantastic portraits
10
and I’m head over heels
In love with you, DNA,
bewitched by your billions
coiled in my cells, transcribing,
replicating, mutating.
I see your never-ending dance.
A length of twisted ladder
briefly unwinds,
both strands duplicate,
each copy drifts away
on its secret mission
to make a thought, feel sunshine,
or digest this morning’s porridge.
Two winding parallel threads,
a tiny tangle of gossamer
designing my life.
DNA, you are astonishing
and I am yours truly.
—Winifred Kavalieris
11
PREFACE TO THE FIFTH EDITION
Chromosomal disorders have been, and will always be, with us; that is a
given. What is changing is our ability to recognize and detect them:
detection both in terms of the subtlety of abnormalities and of the means
we can use to find them. Classical cytogenetics has now well and truly
given way to “molecular karyotyping,” and this has been the extraordinary
development of the early twenty-first century. Readers will now be as
accustomed to molecular nomenclature in defining a segment, such as
chr5:1-18,500,000, as they had been to the classical description,
5p14.1→pter.
The very small deletions and duplications which molecular karyotyping
can now reveal have become familiar to the clinicians and counselors who
see patients and families in the clinic. A large number of these are now on
record, many attracting the nomenclature “copy number variant”: Some
are very well understood, others becoming so, and yet quite a few—
variants of uncertain significance, the acronym “VOUS” in daily parlance
—whose roles in human pathology are imperfectly appreciated. Many are
not in the same mold as the deletions and duplications of classical
cytogenetics, in which the single defect sufficed to cause a particular
phenotype, and always did so: We now need to take account of the concept
of incomplete penetrance, with some microdeletions or duplications not, of
themselves, always leading to an abnormal phenotype. Apparently
clinically normal parents may carry the same alteration as their child with
an abnormal phenotype. Digenic, or “two-hit,” mechanisms may now
require consideration. These were not formerly notions much entering into
the assessment of chromosomal disorders; discussion apropos in the clinic
presents a new challenge.
The number of “new” del/dup syndromes increases almost with each
issue of the clinical genetic journals. We include a mention of a
considerable number of these here (Chapter 14), not intending to create an
encyclopedic resource per se but believing that such a record may provide
a useful first point of contact when these cases are encountered in the
clinic. Copy number variants of uncertain significance, on the other hand,
we mostly take only a broad rather than a detailed view (Chapter 17); the
12
reader will need to consult other repositories for fuller information, as their
interpretations evolve.
The new (or now, established) laboratory methodologies blur the
boundaries between what might have been regarded as the classic
chromosomal abnormalities and Mendelian conditions. Some disorders
recorded as being due not only to segmental deletion/duplication affecting
a single locus but also to point mutation at that locus we continue to treat
as “chromosomal”; and for most, their place in this book is secure. But one
major category, the fragile X syndromes, are now seen as essentially
Mendelian disorders, their historic cytogenetic-based nomenclature
notwithstanding, and they no longer claim their chapter.
Peripheral blood and skin have been the tissues in common usage for
chromosome analysis, with an increasing role for cells got from the
convenient and painless “spit sample.” Prenatal diagnosis has been based
on amniocentesis and chorionic villus sampling, but latterly blastomeres
from early embryos, and fetal DNA in the maternal circulation, have
become targets for testing. Now we can anticipate the potential for whole
genome analysis to be applied to the prenatal diagnosis of the classic
aneuploidies, from a simple maternal blood sample, and this would widen
such testing very considerably. Questions such as these raise ethical issues,
and a literature on “chromosomal ethics” is accumulating.
As we have previously written, however marvelous may be these new
ways to test for chromosomes, the concerns of families remain essentially
the same. We may reproduce here the final paragraph of the Preface of the
first edition of this book, from 1989, as valid now as then:
Families pursue genetic counseling in an effort to demystify the mysterious.
If they did not want to “hear it all,” they would not bother with genetic
counseling. Families want an honest evaluation of what is known and what is
unknown, a clear explanation of all possibilities, both good and bad, and a
sensitive exploration of all available information with which they can make
knowledgeable decisions about future family planning. Thus, Bloch et al.
(1979) succinctly convey the essence of why people go to the genetic
counselor. We hope this book will assist counselors in their task.
Dunedin R.J.M.G.
Melbourne D.J.A.
February 2018
13
ACKNOWLEDGMENTS
14
CONTENTS
15
17. Normal Chromosomal Variation
APPENDICES
A. Ideograms of Human Chromosomes, and Haploid Autosomal
Lengths
B. Cytogenetic Abbreviations and Nomenclature
C. Determining 95 Percent Confidence Limits, and the Standard
Error
References
Index
16
PART ONE
BASIC CONCEPTS
17
1
ELEMENTS OF MEDICAL
CYTOGENETICS
18
whether true or computer-generated false colors. The images produced by
this kaleidoscopic karyotyping could be rather beautiful. Black-and-white
photographs were less splendid but often sufficed (Figure 1–1). Albeit that
molecular methodologies have substantially taken over from classical
cytogenetics, and providing a different view of the genetic material, the
word chromosome will surely last forever.
Chromosomal Morphology
Chromosomes have a linear appearance: two arms that are continuous at
the centromere. Reflecting the French influence in the establishment of the
cytogenetic nomenclature, the shorter arm is designated p (for petit), and
the longer is q (variously explained as being the next letter in the alphabet,
a mistyping of g (for grand), for queue, or as the other letter in the formula
19
p + q = 1). In the early part of the cell cycle, each chromosome is present
as a single structure, a chromatid, a single DNA molecule. During the cell
cycle, the chromosomes replicate, and two sister chromatids form. Now
the chromosome exists as a double-chromatid entity. Each chromatid
contains exactly the same genetic material. This replication is in
preparation for cell division so that, after the chromosome has separated
into its two component chromatids, each daughter cell receives the full
amount of genetic material. It is during mitosis that the chromosomes
contract and become readily distinguishable on light microscopy.
Blood and buccal mucosal cells are the tissues from which DNA is
extracted in routine chromosome analysis. From blood, the nucleated
white cell is the tested component for microarray analysis, and in classical
cytogenetic analysis, it is the lymphocyte. Buccal mucosal cells and white
blood cells2 are obtained from a saliva sample. The chromosomal status of
each small sample is taken as representative of the constitution of
(essentially) every other cell of the body. In the case of invasive prenatal
diagnosis, the cells from amniotic fluid or chorionic villi are the source
material; these tissues are assumed (with certain caveats) to represent the
fetal chromosomal constitution. Noninvasive prenatal diagnosis exploits
the presence of fetal blood cells and DNA in the maternal circulation.
The 46 chromosomes come in 23 matching pairs and constitute the
genome. One of each pair came from the mother, and one from the father.
For 22 of the chromosome pairs, each member (each homolog) has the
same morphology in each sex: These are the autosomes. The sex
chromosome (or gonosome) constitution differs: The female has a pair of
X chromosomes, and the male has an X and a Y chromosome. The single
set of homologs—one of each autosome plus one sex chromosome—is the
haploid set. The haploid number (n) is 23. The haploid complement exists,
as such, only in the gametocytes (ovum and sperm). All other cells in the
body—the soma—have a double set: the diploid complement (2n) of 46. If
there is a difference between a pair of homologs, in the sense of one being
structurally rearranged, the person is described as a heterozygote.
The chromosomes are classically distinguishable on the basis of their
size, centromere position, and banding pattern. The centromere may be in
the middle, off-center, or close to one end—metacentric, submetacentric,
and acrocentric, respectively. The chromosomes are numbered 1 through
22, and X and Y, and are also assigned to groups A through G, according
to their general size and the position of the centromere. The diagrammatic
representation of the banding pattern is the ideogram (Appendix A). The
numbering is based on size, largest to smallest (to split hairs, this order is
20
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Hobrok of hawks, | and Garm of hounds.
[103]
[105]
[106]
King Geirröth sat and had his sword on his knee, half
drawn from its sheath. But when he heard that Othin
was come thither, then he rose up and sought to take
Othin from the fire. The sword slipped from his hand,
and fell with the hilt down. The king stumbled and fell
forward, and the sword pierced him through, and
slew him. Then Othin vanished, but Agnar long ruled
there as king. [84]
[Contents]
NOTES
Prose. The texts of the two manuscripts differ in many minor details.
Hrauthung: this mythical king is not mentioned elsewhere. Geirröth:
the manuscripts spell his name in various ways. [86]Frigg: Othin’s
wife. She and Othin nearly always disagreed in some such way as
the one outlined in this story. Hlithskjolf (“Gate-Shelf”): Othin’s watch-
tower in heaven, whence he can overlook all the nine worlds; cf.
Skirnismol, introductory prose. Grimnir: “the Hooded One.” [87]
2. In the original lines 2 and 4 are both too long for the meter, and
thus the true form of the stanza is doubtful. For line 4 both
manuscripts have “the land of the Goths” instead of simply “the
Goths.” The word “Goths” apparently was applied indiscriminately to
any South-Germanic people, including the Burgundians as well as
the actual Goths, and thus here has no specific application; cf.
Gripisspo, 35 and note. [88]
5. Ydalir (“Yew-Dales”): the home of Ull, the archer among the gods,
a son of Thor’s wife, Sif, by another marriage. The wood of the yew-
tree was used for bows in the North just as it was long afterwards in
England. Alfheim: the home of the elves. Freyr: cf. Skirnismol,
introductory prose and note. Tooth-gift: the custom of making a
present to a child when it cuts its first tooth is, according to
Vigfusson, still in vogue in Iceland.
17. Vithi: this land is not mentioned elsewhere. Vithar avenged his
father, Othin, by slaying the wolf Fenrir. [92]
18. Stanzas 18–20 appear also in Snorri’s Edda. Very possibly they
are an interpolation here. Eldhrimnir (“Sooty with Fire”): the great
kettle in Valhall, wherein the gods’ cook, Andhrimnir (“The Sooty-
Faced”) daily cooks the flesh of the boar Sæhrimnir (“The
Blackened”). His flesh suffices for all the heroes there gathered, and
each evening he becomes whole again, to be cooked the next
morning.
19. Freki (“The Greedy”) and Geri (“The Ravenous”): the two wolves
who sit by Othin’s side at the feast, and to whom he gives all the
food set before him, since wine is food and drink alike for him.
Heerfather: Othin.
20. Mithgarth (“The Middle Home”): the earth. Hugin (“Thought”) and
Munin (“Memory”): the two ravens who sit on Othin’s shoulders, and
fly forth daily to bring him news of the world. [93]
21. Thund (“The Swollen” or “The Roaring”): the river surrounding
Valhall. Thjothvitnir’s fish: presumably the sun, which was caught by
the wolf Skoll (cf. Voluspo, 40), Thjothvitnir meaning “the mighty
wolf.” Such a phrase, characteristic of all Skaldic poetry, is rather
rare in the Edda. The last two lines refer to the attack on Valhall by
the people of Hel; cf. Voluspo, 51.
23. This and the following stanza stand in reversed order in Regius.
Snorri quotes stanza 23 as a proof of the vast size of Valhall. The
last two lines refer to the final battle with Fenrir and the other
enemies.
25. The first line in the original is, as indicated in the translation, too
long, and various attempts to amend it have been made. Heithrun:
the she-goat who lives on the twigs of the tree Lærath (presumably
the ash Yggdrasil), and daily gives mead which, like the boar’s flesh,
suffices for all the heroes in Valhall. In Snorri’s Edda Gangleri
foolishly asks whether the heroes drink water, whereto Har replies,
“Do you imagine that Othin invites kings and earls and other noble
men, and then gives them water to drink?”
31. The first of these roots is the one referred to in stanza 26; the
second in stanza 29 (cf. notes). Of the third root there is nothing
noteworthy recorded. After this stanza it is more than possible that
one has been lost, paraphrased in the prose of Snorri’s Edda thus:
“An eagle sits in the branches of the ash-tree, and he is very wise;
and between his eyes sits the hawk who is called Vethrfolnir.”
33. Stanzas 33–34 may well be interpolated, and are certainly in bad
shape in the Mss. Bugge points out that they are probably of later
origin than those surrounding them. Snorri [98]closely paraphrases
stanza 33, but without elaboration, and nothing further is known of
the four harts. It may be guessed, however, that they are a late
multiplication of the single hart mentioned in stanza 26, just as the
list of dragons in stanza 34 seems to have been expanded out of
Nithhogg, the only authentic dragon under the root of the ash.
Highest twigs: a guess; the Mss. words are baffling. Something has
apparently been lost from lines 3–4, but there is no clue as to its
nature.
36. Snorri quotes this list of the Valkyries, concerning whom cf.
Voluspo, 31 and note, where a different list of names is given. Hrist:
“Shaker.” Mist: “Mist.” Skeggjold: “Ax-Time.” Skogul: “Raging” (?).
Hild: “Warrior.” Thruth: “Might.” Hlokk: “Shrieking.” Herfjotur: “Host-
Fetter.” Gol: “Screaming.” Geironul: “Spear-Bearer.” Randgrith:
“Shield-Bearer.” Rathgrith: Gering guesses “Plan-Destroyer.”
Reginleif: “Gods’-Kin.” Manuscripts and editions vary greatly in the
spelling of these names, and hence in their significance.
38. Svalin (“The Cooling”): the only other reference to this shield is in
Sigrdrifumol, 15.
39. Skoll and Hati: the wolves that devour respectively the sun and
moon. The latter is the son of Hrothvitnir (“The Mighty Wolf,” i.e.
Fenrir); cf. Voluspo, 40, and Vafthruthnismol, 46–47, in which Fenrir
appears as the thief. Ironwood: a conjectural emendation of an
obscure phrase; cf. Voluspo, 40.
40. This and the following stanza are quoted by Snorri. They seem to
have come from a different source from the others of this poem;
Edzardi suggests an older version of the Vafthruthnismol. This
stanza is closely parallel to Vafthruthnismol, 21, which see, as also
Voluspo, 3. Snorri, following this account, has a few details to add.
The stones were made out of Ymir’s teeth and such of his bones as
were broken. Mithgarth was a mountain-wall made out of Ymir’s
eyebrows, and set around the earth because of the enmity of the
giants. [101]
42. With this stanza Othin gets back to his immediate situation,
bound as he is between two fires. He calls down a blessing on the
man who will reach into the fire and pull aside the great kettle which,
in Icelandic houses, hung directly under the smoke-vent in the roof,
and thus kept any one above from looking down into the interior. On
Ull, the archer-god, cf. stanza 5 and note. He is specified here
apparently for no better reason than that his name fits the initial-
rhyme.
43. This and the following stanza are certainly interpolated, for they
have nothing to do with the context, and stanza 45 continues the
dramatic conclusion of the poem begun in stanza 42. This stanza is
quoted by Snorri. Ivaldi (“The Mighty”): he is known only as the
father of the craftsmen-dwarfs who made not only the ship
Skithblathnir, but also Othin’s spear Gungnir, and the golden hair for
Thor’s wife, Sif, after Loki had maliciously cut her own hair off.
Skithblathnir: this ship (“Wooden-Bladed”) always had a fair wind,
whenever the sail was set; it could be folded up at will and put in the
pocket. Freyr: concerning him and his father, see Voluspo, 21, note,
and Skirnismol, introductory prose and note. [102]
45. With this stanza the narrative current of the poem is resumed.
Ægir: the sea-god; cf. Lokasenna, introductory prose. [103]
51. Again the poem returns to the direct action, Othin addressing the
terrified Geirröth. The manuscripts show no lacuna. Some editors
supply a second line from paper manuscripts: “Greatly by me art
beguiled.”
53. Ygg: Othin (“The Terrible”). The maids: the three Norns.
54. Possibly out of place, and probably more or less corrupt. Thund:
“The Thunderer.” Vak: “The Wakeful.” Skilfing: “The Shaker.” Vofuth:
“The Wanderer.” Hroptatyr: “Crier of the Gods.” Gaut: “Father.” Ofnir
and Svafnir: cf. stanza 34. [107]
[Contents]
SKIRNISMOL
The Ballad of Skirnir
[Contents]
Introductory Note
The Skirnismol is found complete in the Codex Regius, and through
stanza 27 in the Arnamagnæan Codex. Snorri quotes the concluding
stanza. In Regius the poem is entitled “For Scirnis” (“Skirnir’s
Journey”).
The Skirnismol differs sharply from the poems preceding it, in that it
has a distinctly ballad quality. As a matter of fact, however, its verse
is altogether dialogue, the narrative being supplied in the prose
“links,” concerning which cf. introductory note to the Grimnismol. The
dramatic effectiveness and vivid characterization of the poem seem
to connect it with the Thrymskvitha, and the two may possibly have
been put into their present form by the same man. Bugge’s guess
that the Skirnismol was the work of the author of the Lokasenna is
also possible, though it has less to support it.
[Contents]
Skirnir spake:
[109]
Skirnir spake:
Freyr spake:
Skirnir spake:
Freyr spake:
[110]
7. “To me more dear | than in days of old
Was ever maiden to man;
But no one of gods | or elves will grant
That we both together should be.”
Skirnir spake:
Freyr spake:
[111]
Skirnir spake:
Gerth spake: