DEPARTURES FROM DIPLOIDY
##
  The diploid chromosome complement of Drosophila
  melanogaster may be designated X/X; 2/2;
  3/3; 4/4 for females and X/Y; 2/2; 3/3;
  4/4 for males. Addition to or subtraction from
  either of these complements of one or more
  chromosomes produces a departure from diploidy. The
  non-diploid constitutions are designated by a name but
  not a symbol except as included in the name, e.g.,
  X/0 male. Constitutions are described by listing
  their component chromosomes, homologous chromosomes
  being separated by slash bars and nonhomologous
  chromosomes by semicolons. When two homologous
  chromosomes are attached to the same centromere,
  components are listed without separation, e.g., XX,
  XY, and 44.
#
diploid metafemale
 constitution: X/X/X; 2/2; 3/3; 4/4; sex chromosome
  constitution may also be XX/X.
 origin: Produced by triploid and compound-X-bearing
  females. May result from two-X gametes produced
  by nondisjunction.
 discoverer: Bridges.
 synonym:superfemale.
 references: 1921, Science 54: 252-54.
  1922, Am. Naturalist 56: 51-63 (fig.).
  1925, Am. Naturalist 59: 127-37.
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 153-62 (fig.).
 phenotype: :Wings crumpled or incised on inner margin. Rear
  legs often malformed. Viability low, usually less
  than 0.5%. Flies die mostly in late larval and pupal
  stages; at 25[o]C, puparium formation delayed 1-2 days
  (Brehme, 1937,
  Proc. Soc. Exptl. Biol. Med. 37: 578-80).
  Survivors sterile; two fertile metafemales
  were apparently mosaic for triploid tissue [Rolfes
  and Hollander, 1961, J. Heredity 52: 61-66
  (fig.)]. Larval ovaries from metafemales
  transplanted into sterile diploids have produced a
  few progeny (Beadle and Ephrussi, 1937,
  Proc. Natl. Acad. Sci. U.S. 23: 356-60). Crossing over between
  the X chromosomes appears to be infrequent.
 other information: The term metafemale instead of superfemale was
  suggested by Stern (1959, Lancet 12: 1088).
#
haplo-4
 constitution: X/X; 2/2; 3/3; 4; sex chromosome constitution may
  also be X/Y.
 origin: Produced after occasional loss or nondisjunction of
  chromosome 4 during meiosis. Produced in
  quantity from crosses of C(4)RM/0 with normal
  or from heterozygous T(2;4) or T(3;4)
  females.
 discoverer: Bridges, 20a30.
 references: 1921, Proc. Natl. Acad. Sci. U.S. 7: 186-92.
  1922, Am. Naturalist 56: 51-63 (fig.).
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 135-43 (fig.).
 phenotype: Minute phenotype:  caused by deficiency for M(4).
  Pale body with prominent trident pattern on thorax.
  L5 often does not reach wing margin. Eclosion
  delayed 2-4 days. Viability erratic, usually below
  80% of normal. Usually sterile. Male tends to be
  more viable and fertile than female.
#
haploid
 constitution: X; 2; 3; 4;.
 origin: Recorded as patches of tissue.
 discoverer: Bridges.
 references: 1925, Proc. Natl. Acad. Sci. U.S. 11: 706-10.
  1930, Science 72: 405-6.
 phenotype: Eye facets small in haploid patches. A haploid
  foreleg bore no sex comb; the tissue is therefore
  probably female, as expected on the basis of balance
  theory of sex determination.
#
intersex
 constitution: X/X; 2/2/2; 3/3/3; 4/4/4; presence of Y and
  number of fourth chromosomes variable.
 origin: Regularly found among progeny of triploid females.
 discoverer: Bridges, 20l.
 references: 1921, Science 54: 252-54.
  1922, Am. Naturalist 56: 51-63 (fig.).
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 153-62 (fig.).
  Bridges, 1939, In Sex and Internal Secretions
  (E. Allen, C. H. Danforth, and C. A. Doisy, eds.). The
  Williams and Wilkins Co., pp. 15-63.
 phenotype: Large-bodied fly with coarse bristles, roughish
  eyes, and scalloped wing margins. Small hairs on
  surface of wing more sparsely distributed than in
  diploids. Usually has sex combs and a mixture of
  male and female genitalia; genitalia may be malelike
  or femalelike. Addition of sections of X
  chromosome shifts intersexes toward femaleness
  [Dobzhansky and Schultz, 1934, J. Genet. 28: 349-86
  (fig.); Pipkin, 1940, Univ. Texas Publ. 4032: 126-56].
  Addition of sections of the second or the
  third chromosome has not resulted in a shift of
  sexuality (Pipkin, 1947, Genetics 32: 592-607;
  1960, Genetics 45: 1205-16). Fung and Gowen
  reported that a triploid line producing intersexes
  with preponderantly female genitalia carries several
  fourth chromosomes and that another triploid line
  producing male-like intersexes carries only two fourth
  chromosomes.
#
metamale
 constitution: X/Y; 2/2/2; 3/3/3; 4/4/4; inferred from markers
  inherited. May also be diplo-4.
 origin: Occurs among progeny of triploid female.
 discoverer: Bridges, 20l.
 synonym:supermale.
 references: 1921, Science 54: 252-54.
  1922, Am. Naturalist 56: 51-63 (fig.).
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 153-62 (fig.).
 phenotype: Male has small body and spread wings. Late
  hatching, poorly viable, and completely sterile.
#
nullo-X
 constitution: Y/Y; 2/2; 3/3; 4/4.
 origin: One-fourth the progeny from crosses between certain
  compound-X-bearing females (e.g.,
  C(1)RM/Y) and normal males.
 phenotype: Dies as embryo (Li, 1927, Genetics 12: 1-58).
  Cleavage nuclei abnormally distributed and
  blastoderm not formed (Poulson, 1940,
  J. Exptl. Zool. 83: 271-325). According to Scriba
  (1964, Zool. Jahrb. Abt. Anat. Ontog. Tiere 81: 435-90),
  migration of cleavage nuclei to surface of
  egg is normal, blastoderm formation irregular, and germ
  band development frequently incomplete.
#
nullo-X nullo-Y
 constitution: 2/2; 3/3; 4/4.
 origin: One-fourth the progeny of crosses such as
  C(1)RM/0 females with
  Y[S]X.Y[L]/0 males.
 phenotype: Most embryos die after 10-12 cleavages
  (von Borstel and Rekemeyer, 1958, Nature 181: 1597-98).
  Embryology like that of nullo-X (Scriba, 1964,
  Zool. Jahrb. Abt. Anat. Ontog. Tiere 81: 435-90).
#
tetra-4
 constitution: X/X; 2/2; 3/3; 4/4/4/4. Sex chromosome
  constitution may also be X/Y; that for
  chromosome 4 may be 44/4/4 or 44/44.
 references: Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 135-43.
  Li, 1927, Genetics 12: 1-58.
  Bridges, 1935, Tr. Dinam. Razvit. 10: 463-74.
  Grell, 1961, Genetics 46: 1177-83 (fig.).
 phenotype: Viability reduced; usually dies in embryonic or
  larval stage. Wings of survivors longer and more
  pointed than normal.
 origin: Synthesized as females homozygous for T(1;4)w[m4]
  + T(1;4)B[S] formed by recombination in region
  3C4-15F8 between T(1;4)w[m5] =
  T(1;4)3C3-4;101F1-2 and T(1;4)B[S] =
  T(1;4)15F9-16A1;16A7-B1;102F (Grell, 1961). Also
  recovered from progeny of crosses between males and
  females that carry C(4)RM (E. B. Lewis).
#
tetraploid
 constitution: X/X/X/X; 2/2/2/2; 3/3/3/3; 4/4/4/4.
 origin: Seen on a few occasions as a tetraploid daughter of
  a triploid female or as a patch of tetraploid gonial
  tissue in an otherwise diploid female. Extensive
  attempts to produce tetraploid males have failed.
 discoverer: Bridges.
 references: 1925, Am. Naturalist 59: 127-37.
  Morgan, 1925, Genetics 10: 148-78.
 phenotype: Recognized by production of progeny that are almost
  exclusively triploids and intersexes.
#
triplo-4
 constitution: X/X; 2/2; 3/3; 4/4/4. Sex chromosome constitution
  may be X/Y; that for chromosome 4 may be
  44/4.
 origin: Product of nondisjunction of chromosome 4.
  Regular product of cross between C(4)RM and
  normal diplo-4 flies.
 discoverer: Bridges, 21b13.
 references: 1922, Am. Naturalist 56: 51-63.
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 21 (fig.), 135-43.
 phenotype: Phenotypic departure from normal very slight. Body
  darker than normal and trident pattern subdued. Eyes
  small. Body and wings elongate. Preferential
  segregation of the different fourth chromosomes in
  triplo-4's described by Sturtevant (1936,
  Genetics 21: 444-66).
#
triploid
 constitution: X/X/X; 2/2/2; 3/3/3; 4/4/4. Sex chromosome
  constitution may also be X/X/X/Y, XX/X, or
  XX/X/Y. Triploids from stocks kept for several
  generations usually carry only two fourth
  chromosomes, i.e., diplo-4 triploids.
 origin: Spontaneous from unreduced eggs; incidence increased
  by treatment with cold (Bauer, 1946,
  Z. Naturforsch. 1: 35-38; Gloor, 1950, DIS 24: 82) or with
  colchicine (Braungart and Ott, 1942, Sci. Counselor
  8: 105; Schultz). Produced in relatively high
  frequency by triploid females and by c(3)G/c(3)G
  females (Gowen, 1933, J. Exptl. Zool. 65: 83-106).
 discoverer: Bridges, 1920.
 references: 1921, Science 54: 252-54.
  1922, Am. Naturalist 56: 51-63 (fig.).
  Morgan, Bridges, and Sturtevant, 1925,
  Bibliog. Genet. 2: 135-43.
 phenotype: Eye facets larger and hairs on surface of wings more
  sparsely distributed than in diploid, giving eyes and
  wings a coarse texture; bristles also coarse. These
  characteristics are diagnostic for three sets of
  autosomes and result from increased cell size. Body
  thickset. Ventral bristles between first and second
  pairs of legs often missing. Discernible from
  diploid with practice. Fertility poor owing to
  production of aneuploid classes of gametes. Because
  equal numbers of chromosomes tend to go to each pole
  during first meiotic division, euploid
  gametes are produced with lower-than-expected
  frequencies; gametes with one X and two
  sets of autosomes and with two X's and one set
  of autosomes far outnumber those with one X and one
  set of autosomes or two X's and two sets of
  autosomes (Bridges and Anderson, 1926, Genetics
  10: 418-41). Triploids that carry an attached X
  (attached X triploids) are more fertile and
  produce a higher proportion of triploid progeny than
  free X triploids. Triploids are of necessity
  female and their progeny include metafemales,
  metamales, intersexes, triploid and diploid females,
  and diploid males. Crossing over is markedly increased
  in triploids; Sturtevant (1951,
  Proc. Natl. Acad. Sci. U.S. 37: 405-7) has mapped
  chromosome 4 in
  diplo-4 triploids. B, Bl, Bx, Cy, D, Dfd, H,
  Hw, J, L[2], Me, and Sb are classifiable in a
  single dose in triploids. Dl, G, N, bw[V1], Px,
  S, and all Minutes are recessive in a single dose.
  Two doses of D, Dl, G, H, bw[V1], Px, and Sb
  produce an extreme phenotype: , whereas two doses of
  M or Me are lethal (Schultz, 1934, DIS 1: 55).
#
triploid metafemale
 constitution: X/X/X/X; 2/2/2; 3/3/3; 4/4/4; third 4 may be
  absent.
 origin: Found among progeny of tetraploid female (Morgan).
  Also produced by nondisjunction of sex chromosomes
  in C(1)RM/In(1)sc[8]/Y triploid (Frost).
 discoverer: L. V. Morgan.
 references: 1925, Genetics 10: 147-78.
  Frost, 1960, Proc. Natl. Acad. Sci. U.S. 46: 47-51.
 phenotype: Coarse eyes, wing texture, and bristles. Resembles
  triploid except body smaller and eyes more bulging.
  Inner wing margins often incised. Using exceptional
  triploid females as a standard, Frost (1960)
  determined that triploid metafemales have 25%
  viability, 24-54% lay eggs (1 to 150 eggs),
  and about 11% of the eggs develop into adults.
#
X0 male
 constitution: X; 2/2; 3/3; 4/4.
 origin: Product of primary nondisjunction of the sex
  chromosomes in either father or mother in cross of
  X/Y male with X/X female. Forms one-fourth
  the progeny of crosses, such as X/X female by
  Y[S]X.Y[L]/0 male or
  C(1)RM/0 female by X/Y male.
 discoverer: Bridges.
 references: 1916, Genetics 1: 1-52.
 phenotype: Male morphologically normal but entirely sterile.
  No motile sperm produced. Primary spermatocyte
  nuclei lack the morphological elements
  characteristic of normal male; these elements
  replaced by needle-shaped crystals that are found
  in the nucleus, the cytoplasm, and extracellularly
  (Meyer, Hess, and Beermann, 1961, Chromosoma 12: 676-716).
  Nebenkern and axial filament differentiation
  during spermiogenesis is abnormal (Kiefer, 1966,
  Genetics 54: 1441-52).
#
XXY female
 constitution: X/X/Y; 2/2; 3/3; 4/4. Sex chromosome
  constitution may also be X/XY or XX/Y.
 origin: Product of either primary or secondary
  nondisjunction in either male or female. Also
  produced from cross of an XY-bearing parent with
  a normal-X-bearing parent. Condition usually
  found in compound-X-bearing female.
 discoverer: Bridges.
 references: 1916, Genetics 1: 1-52.
 phenotype: Phenotype and fertility like those of normal female.
  Nondisjunction of X chromosomes in X/X/Y
  much higher than in X/X female; about 4%
  exceptions with two normal X chromosomes and much
  higher if X's are heterozygous for inversions
  (Sturtevant and Beadle, 1936, Genetics 21: 554-604).
#
XXYY female
 constitution: X/X/Y/Y; 2/2; 3/3; 4/4. Sex constitution
  may also be XX/Y/Y, X/XY/Y, or XY/XY.
 origin: A common product of crosses such as
  Y[S]X.Y[L]/Y male by
  X/Y[S]X.Y[L]
  or X/X/Y female, or X/Y/Y male by
  X[S]X.Y[L]/X, X/X/Y, or
  C(1)RM/Y female.
 discoverer: Stern.
 references: 1929, Biol. Zentr. 49: 261-90; 727.
  Cooper, 1956, Genetics 41: 242-64.
 phenotype: Eye color mottled to varying degrees. Posterior and
  middle legs often malformed. Fertility and
  viability reduced. Gametes preponderantly X/Y
  in constitution owing to the regular segregation of
  both the X's and the Y's at the first
  meiotic division.
#
XYY male
 constitution: X/Y/Y; 2/2; 3/3; 4/4. Sex chromosome consitution
  may aso be XY/Y.
 origin: About one-fourth the progeny of crosses such as
  X/X/Y female by X/Y male, C(1)RM/Y
  female by Y[S]X.Y[L]/Y male,
  and X/Y[S]X.Y[L] female by
  X/Y male.
 discoverer: Bridges.
 references: 1916, Genetics 1: 1-52.
 phenotype: Phenotype normal; usually fertile; with certain
  normal Y chromosomes, completely sterile
  (R. F. Grell). The two Y chromosomes tend to
  separate at the first meiotic division to a degree
  depending on the source of the Y's and the
  X (Grell, 1958,
  Proc. Intern. Congr. Genet. 10th., Vol. 2: 105).
#
XYYY male
 constitution: X/Y/Y/Y; 2/2; 3/3; 4/4. Sex chromosome
  constitution may also be XY/Y/Y.
 discoverer: Stern.
 references: 1929, Biol. Zentr. 49: 261-90.
  Morgan, Bridges, and Schultz, 1934, Carnegie
  Inst. Wash. Year Book 33: 274-80.
  Cooper, 1956, Genetics 41: 242-64.
 phenotype: Morphologically normal male but with mottled eyes
  as in XXYY female. Almost entirely sterile;
  Cooper (1956) suggests that the few offspring may
  result from X/Y/Y cysts produced by mitotic
  loss of a Y chromosome.
#