Euphytica 104: 95–106, 1998.

© 1998 Kluwer Academic Publishers. Printed in the Netherlands.

95

Plant aneuploids: Suggestions for their classification

A.K. Koul & M.K. Dhar∗

Department of Biosciences, University of Jammu, Jammu-180006, India; (∗ author for correspondence)

Received: 25 March 1997; accepted 5 June 1998

Key words: aneuploids, nomenclature, trisomics, monosomics, gene content

Summary
The paper lists, defines and classifies all types of aneuploids known hitherto. Categorisation has been made on the
basis of the alterations of chromosome number and gene content. The change in nomenclature of some aneuploids
has been warranted by the fact that some names that are in vogue do not reflect the precise nature of the aneuploid
and, are therefore, ambiguous. For these ambiguous terms new names have been proposed.

Introduction loss or gain involved even a chromosome segment.

In this way, segmental aneuploidy was added to the
general category of aneuploids. Segmental aneuploidy
Aneuploidy refers to the nuclear condition when the
represents heterozygosity for such deletions or dupli-
chromosome count is not exact multiple of the base
cations whereby one or more acentric segments of
number of the species. The term was introduced by
a basic set are added or lost. It also includes those
Tackholm (1922) following his studies on genus Rosa.
centric fragments where new chromosome arm com-
Aneuploids generally arise through the fusion of ga-
binations have been added (modified from Rieger et
metes, one or both of which are hypo- or hyperhaploid.
al., 1976).
Such gametes arise through anomalies in the meiotic
The classification of aneuploids proposed by ear-
division accompanying gametogenesis. The anomaly
lier workers revolves around the nature of change in
most significant in terms of the differentiation of ga-
chromosome number. In some cases, aneuploidy does
metes with deviant numbers is, late disjunction or
not change the total chromosome count of the nucleus,
non-disjunction of chromosomes during gametogene-
but alters the number of specific chromosomes, the
sis.
total remaining the same. Khush (1973) identified 31
Lutz (1909) was the first to report aneuploid chro-
kinds of aneuploids. Since then, many more types have
mosome numbers in two progenies of the interspecific
been discovered; the number has grown to 74.
cross between Oenothera lata and O. gigas. After the
The present paper aims at listing all aneuploids
discovery of the ‘Globe’ mutant in Datura stramonium
known hitherto at one place and classifying them on
by Avery in 1915, Blakeslee et al. (1920) produced
the basis of their chromosome constitution and gene
various other aneuploids in this species. Realization
content. Changes in chromosome number and gene
of the possible utility of aneuploids generated a spurt
dosage may or may not correspond. For example,
of activity, and as a consequence much literature has
all secondary trisomics carry an isochromosome in
accumulated on the subject in the last 75 years (see
addition to the standard complement. In such indi-
Burnham, 1962; Khush, 1973; Bond & Chandley,
viduals the chromosome number increases only by
1983; Dellarco et al., 1985; Koul et al., 1995).
one, though the genes located on the isochromosome
The term aneuploid was initially used for indi-
are actually represented four times. Should such indi-
viduals with one chromosome less or more than the
viduals be classified as tri- or tetrasomic? Similarly,
standard complement. The former were named tri-
individuals trisomic for one and monosomic for an-
somics and the latter monosomics. Dyer et al. (1970)
other chromosome do not register any change in total
extended the term to cells and individuals where the

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Table 1. Abbreviations used in designating aneuploids
Abbreviation Definition
Tri Trisomic
p primary
s secondary
t telo; telocentric chromosome
te tertiary
a acrocentric chromosome
i isochromosome
I Inversion
(I) Interchange
chromosome number. Yet they differ from the disomic
with respect to dosage of different genes. The names
proposed for such aneuploids by earlier workers do not
reflect their true nature. We have made an attempt to
do away these and other ambiguities existing in the
nomenclature of aneuploids by classifying them on the
basis of changes in chromosome number as well as
gene dosage.
Proposed classification
The 74 aneuploid types known hitherto have been
grouped into five categories. Each category includes
one or more subtypes. Each aneuploid type has been
given an abbreviation and a chromosomal formula.
The formulae are based on the tomato system (Khush
& Rick, 1967). In all formulae, 2n refers to the
disomic chromosome count; 2n + 1 and 2n− 1 there-
fore, denote addition and deficiency of a full chro-
mosome, respectively. Individual chromosomes have
been numbered A, B, C — etc. Chromosome arms are
designated AS.AL, BS.BL etc., L denoting the long
and S the short arm. Addition of the whole chromo-
some has been denoted as 2n + AS.AL, 2n + BS.BL,
etc. Isochromosomes for the short and long arms of
chromosome A are represented as AS.AS and AL.AL
respectively. The telocentrics for long and short arms
of chromosome A are represented as .AL and .AS.
The tertiary chromosome involving chromosomes A
and B is represented as AS.BL suggesting that the
interchanged chromosome carries the long arm of
chromosome B and short arm of chromosome A.
Table 1 lists the letters which have been used
singly or in combination for designating the aneuploid
types. The idiograms of all aneuploid types have been
illustrated in Figure 1. The karyotype of Plantago
ovata (2n = 8), was used as an example. It was se-
lected mainly because, the chromosome number of the
species is low, all chromosomes are heterobranchial
and are morphologically distinguishable from one an-
other. Recently Raupp et al. (1995) have proposed a
nomenclature for the genetic stocks of wheat. Sev-
eral conventions regarding the designation of aneu-
ploids have been enumerated and rules of hierarchy
established. These conventions and rules of hierar-
chy have been followed in the present communication,
whereever applicable.
As pointed out before, the 74 known aneuploid
types have been classified into five main categories.
These are listed below:
A. Addition aneuploids
B. Compensation aneuploids
C. Deletion aneuploids
D. Two-in-one aneuploids
E. Pseudo aneuploids
A. Addition aneuploids
This class includes aneuploids which carry in addition
to the standard complement one or more, chromo-
some(s), chromosome arm(s) or parts thereof. The
extra elements exist as acrocentric, telocentric, meta-
centric or isochromosomes. The addition aneuploids
are subdivided into following three types on the basis
of the number of additional chromosomes.
I. Single chromosome additions. Aneuploids of this
category carry one chromosome in addition to the
standard complement. It includes a number of sub-
types which are listed below.
1. Primary trisomic pTri: An individual in which the
additional chromosome is one from the standard
complement (2n + AS.AL).
2. Secondary trisomic sTri: A trisomic in which the
additional chromosome is an isochromosome for
one chromosome arm of the standard complement
(2n + DL.DL).
3. Tertiary trisomic teTri: The additional chromo-
some consists of parts belonging to two non-
homologous chromosomes (tertiary chromosome)
of the standard complement (2n + BS.AL).
4. Telotrisomic tTri: These aneuploids have a telo-
centric arm added to the standard complement
(2n + .DL).
5. Inversion telotrisomic ItTri: The additional ele-
ment in these trisomics is a telocentric chromo-
euph4423.tex; 5/10/1998; 7:06; p.2
 97

Figure 1. Idiograms of different aneuploid types described. Abbreviations: p – primary; s – secondary; te – tertiary; t – telocentric, telosomic;
i – iso, isosomic; a - acro; (I) Interchange; I – Inversion; Tri – trisomic; T – tetrasomic; P – pentasomic; Q – quadruple; H – hexasomic; O –
octosomic; C – compensating; M – monosomic; N – nullisomic; (trp) – triple; D – di, disomic; d – double; (tet) – tetra.

euph4423.tex; 5/10/1998; 7:06; p.3

98

Figure 1. Continued.

euph4423.tex; 5/10/1998; 7:06; p.4


Figure 1. Continued.
some which is the product of pericentric inversion
and misdivision of the centromere. The extra chro-
mosome consists of segments belonging to both
arms of the chromosome (2n + .DSDL ).
6. Metatrisomic mTri: Aneuploid with an additional
metacentric chromosome that is unlike any in the
basic complement, having arisen through struc-
tural alteration of some chromosome of the stan-
dard complement (2n + meta DSDL ).
7. Acrotrisomic aTri: An aneuploid in which the ex-
tra chromosome is acrocentric, having originated
through deletions of terminal end(s) of a chro-
mosome of the standard complement (2n + acro
DSDL ).
8. Interchange primary trisomic (I)pTri: A transloca-
tion heterozygote carrying an extra chromosome
the homologue of which may or may not be in-
volved in the interchange (2n(TAL.BL) + AS.AL).
9. Interchange secondary trisomic (I)sTri: A translo-
cation heterozygote carrying an additional isochro-
99
mosome for one arm of a chromosome involved in
translocation (2n(TAL.BL) + AL.AL).
10. Interchange tertiary trisomic (I)teTri: A translo-
cation heterozygote carrying an additional tertiary
chromosome for those chromosomes involved in
the translocation (2n(TAL.BL) + AS.BS).
From within the above 10 types, 4 types namely pri-
mary, secondary, tertiary and telo trisomics have been
recovered in a number of species and studied in detail
(Khush, 1973).
II. Two chromosome additions. These aneuploids
carry two chromosomes in addition to the standard
complement.
1. Double trisomic dTri: An individual trisomic for
two different chromosomes of the standard com-
plement (2n + AS.AL + DS.DL).
2. Diacrotrisomic DaTri: An aneuploid carrying two
homologous acrocentric chromosomes [result of
break(s) at the end(s)] besides the standard com-
plement. The aneuploid is partial tetrasomic for
euph4423.tex; 5/10/1998; 7:06; p.5
100
a part of one of the chromosomes of the standard
complement (2n + 2 acro DSDL ).
3. Tetrasomic T: An individual having an extra pair
of homologous chromosomes from the standard
complement [2n + 2 (AS.AL)].
4. Monotertiary tetrasomic MteT: An individual hav-
ing a normal and a tertiary chromosome in addi-
tion to the standard complement (2n + AS.AL +
BS.AL).
5. Ditertiary tetrasomic DteT: An individual having
two additional homologous tertiary chromosomes
[2n + 2(BS.AL)].
6. Ditelotetrasomic DtT: An individual having an
additional pair of homologous telocentric chromo-
somes [2n + 2(.DL)].
7. Inversion telotetrasomic ItT: The aneuploid is pri-
mary trisomic and inversion telotrisomic for the
same chromosome (2n + DS.DL + .DSDL ).
8. Inversion ditelotetrasomic IDtT: An individual
having an extra pair of homologous telocentric
chromosomes; the telocentrics being the result
of pericentric inversion and misdivision [2n +
2(.DSDL )].
9. Inversion telotrisomic plus primary trisomic ItTri
pTri: An aneuploid primary trisomic for one chro-
mosome and inversion telotrisomic for another
chromosome of the complement (2n + AS.AL +
.DSDL ).
III. Three or more chromosome additions. These
aneuploids carry three or more chromosomes in ad-
dition to the basic complement.
1. Triple trisomic (trp)Tri: An individual trisomic
for three different chromosomes of the standard
complement (2n + AS.AL + BS.BL + DS.DL).
2. Tetra trisomic (tet)Tri: An individual which is
tetrasomic for one chromosome and trisomic for a
different chromosome [2n + AS.AL + 2(DS.DL)].
3. Inversion ditelotetrasomic plus primary trisomic
IDtT pTri: A primary trisomic carrying in addi-
tion two telocentric chromosomes [2n + AS.AL +
2(.DSDL )].
4. Pentasomic P: An individual carrying three homol-
ogous chromosomes in addition to the standard
complement so that this chromosome is repre-
sented five times [2n + 3(DS.DL)].
5. Inversion tritelopentasomic ItP: An individual hav-
ing three additional homologous telocentric chro-
mosomes. The aneuploid is pentasomic for some
segments [2n + 3(.DSDL)].
6. Inversion tritelopentasomic plus trisomic ItPTri: A
primary trisomic that has in addition three homol-
ogous telocentric chromosomes [2n + AS.AL +
3(.DSDL)].
7. Quadruple trisomic QTri: An aneuploid, trisomic
for four different chromosomes of the standard
complement [2n + AS.AL + BS.BL + CS.CL +
DS.DL).
8. Hexasomic H: Aneuploid in which one chromo-
some of the complement is represented six times
[2n + 4(DS.DL)].
9. Inversion tetratelo hexasomic ItH: An individ-
ual having four additional homologous telocen-
tric chromosomes. The aneuploid is as a result
hexasomic for some segments [2n + 4(.DSDL )].
10. Octosomic O: Aneuploid in which one chromo-
some is represented eight times [2n + 6(AS.AL)].
This category of aneuploids is the least studied. Meio-
sis of these aneuploids is characterized by associations
which reflect their chromosome constitution (Table 1).
B. Compensation aneuploids
This category comprises aneuploids in which loss of
a chromosome of the standard complement is com-
pensated by presence of more than one altered chro-
mosomes, leading to increase in the number of chro-
mosomes or chromosome segments. This class of
aneuploids includes following types.
1. Telo-iso compensating trisomic tiCTri: A trisomic
with the missing chromosome compensated for by
one iso- and one telocentric chromosome (2n –
AS.AL + AS.AS + .AL).
2. Telo-tertiary compensating trisomic tteCTri: A
compensating trisomic with the missing chromo-
some compensated for by one telocentric and
one tertiary chromosome (2n – AS.AL + .AS +
BS.AL).
3. Diiso compensating trisomic DiCTri: A trisomic
from which one chromosome is missing but its
loss is compensated by two isochromosomes, one
for each arm of the missing chromosome (2n –
DS.DL + DS.DS + DL.DL).
4. Ditertiary compensating trisomic DteCTri: A tri-
somic in which the missing chromosome is com-
pensated for by two tertiary chromosomes, one
having long and the other short arm of the missing
chromosome (2n – AS.AL + AS.BL + BS.AL).
5. Iso-tertiary compensating trisomic iteCTri: A tri-
somic in which the missing chromosome is com-
euph4423.tex; 5/10/1998; 7:06; p.6

pensated for by one iso- and one tertiary chromo-
some (2n – AS.AL + AS.AS + BS.AL).
6. Double telotrisomic plus primary trisomic dtTri
pTri: An individual trisomic for one chromosome,
monosomic for the other, and carrying in addition
two telocentrics one for each arm of the missing
chromosome (2n – AS.AL + .AS + .AL + DS.DL).
7. Double telotrisomic plus tertiary trisomic dtTri
teTri: An individual monosomic for one chromo-
some and having instead a tertiary chromosome
and two telocentrics. The latter compensate for
the missing chromosome of the basic complement
(2n – AS.AL + .AS + .AL + BS.DL).
8. Ditelo diiso tetrasomic DtDiT: An individual in
which the loss of two chromosomes is compen-
sated by two telocentrics and two isochromosomes
[2n – 2(AS.AL) + 2(.AL) + 2(AS.AS)].
9. Dimonotelo tetrasomic DMtT: A monosomic with
the missing chromosome compensated for by three
telocentrics, two for one arm and one for the
second arm of the missing chromosome [2n –
AS.AL + 2(.AS) + .AL)].
10. Dimonotelo tetrasomic plus primary trisomic
DMtT pTri: This individual is primary trisomic for
one chromosome, deficient for the other chromo-
some and has in addition three telocentrics, two
for one and one for the other arm of the miss-
ing chromosome [2n – AS.AL + 2(.AS) + .AL +
DS.DL].
11. Dimonotelo tetrasomic plus double telotrisomic
DMtT dtTri: An individual deficient for two non-
homologous chromosomes, having instead five te-
locentrics, two representing an arm of one missing
chromosome and the other three compensating for
the three other missing arms (two of one and the
third of the second chromosome) [2n – AS.AL –
BS.BL + 2(.AS) + .AL + .BS + .BL)].
12. Double ditelopentasomic dDtP: A monosomic in
which the missing chromosome is compensated by
four telocentrics, two for each arm [2n – AS.AL +
2(.AS) + 2(.AL)].
C. Deletion aneuploids
This category includes aneuploids deficient for one or
more chromosomes or chromosome segments. They
are of the following types.
1. Monosomic M: Aneuploids deficient in one chro-
mosome of the standard complement (2n –
DS.DL).
101
2. Tertiary monosomic teM: Aneuploids in which
the arms of two nonhomologous chromosomes are
missing and the remaining two arms have joined
forming a tertiary chromosome (2n – AS.BL).
3. Monotelodisomic MtD: Aneuploids deficient in
one chromosome arm (2n – .AS).
4. Monotelosomic Mt: Aneuploids missing one full
chromosome and one arm of the other homologue
[2n – 2(AS.AL) + .AS].
5. Ditelosomic Dt: An individual missing two homol-
ogous arms [2n – 2(AS.AL) + 2(.AS)].
6. Double monotelosomic dMt: A nullisomic with
two telocentrics representing two arms of one
chromosome of the missing pair [2n – 2(AS.AL) +
.AS + .AL)].
7. Monotelo monoisosomic MtMi: A nullisomic in
which one arm of the missing pair is represented
by a telocentric and the other arm by an isochro-
mosome [2n – 2(AS.AL) + AS.AS + .AL)].
8. Monoisosomic Mi: A nullisomic in which one arm
of the missing pair is represented as an isochromo-
some [2n – 2(AS.AL) + AS.AS)].
9. Ditelo monotelosomic DtMt: An individual from
which one chromosome pair is missing but three
telocentrics, two for one arm and one for another
arm of the missing chromosome are present [2n –
2(AS.AL) + .AS +2(.AL)].
10. Nullisomic N: Aneuploids deficient for one ho-
mologous chromosome pair [2n – 2(AS.AL)].
11. Double monosomic dM: Aneuploids deficient for
two non-homologous chromosomes (2n - AS.AL
– BS.BL).
12. Triple monosomic (trp)M: Individuals from which
three different chromosomes are missing (2n –
AS.AL – BS.BL – CS.CL).
13. Double nullisomic dN: An individual from which
two pairs of chromosomes are missing [2n –
2(AS.AL) – 2(BS.BL)].
From the aneuploids listed above monosomics and
nullisomics have been raised and utilized extensively
in several crop plants especially wheat, oats, and
cotton (Burnham, 1962; Khush, 1973).
D. Two-in-one aneuploids
These individuals represent addition aneuploids for
some chromosome (s)/chromosome part(s) and dele-
tion aneuploids for other chromosome(s)/chromosome
part(s) of the standard set. In these aneuploids the stan-
dard complement is not complete. The following types
are referred to this category.
euph4423.tex; 5/10/1998; 7:06; p.7
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Table 2. Cytological characteristics of various types of aneuploids

Aneuploid Reference Plant species Characteristic meiotic

configuration

A. Addition aneuploids
I. Single chromosome additions
1. pTri Goodspeed & Avery, 1939 Nicotiana sylvestris (n-1) II + 1III
2. sTri Goodspeed & Avery, 1939 Nicotiana sylvestris (n-1) II + 1III
3. teTri Goodspeed & Avery, 1939 Nicotiana sylvestris (n-2) II + 1V
4. tTri Khush & Rick, 1968 Lycopersicon esculentum (n-1) II + 1III
5. ItTri Dhar et al., 1990; Dhar 1991 Plantago lagopus (n-1) II + 1III
6. mTri Fujigaki & Tsuchiya, 1981 Hordeum vulgare (n-1) II + 1III
7. aTri Tsuchiya & Fujigaki, 1981 Hordeum vulgare (n-1) II + 1III
8. (I) pTri Burnham, 1930 Zea mays (n-2) II + 1V
9. (I)sTri Rao et al., 1988 Pennisetum americanum (n-2) II + 1V
10. (I)teTri Rao et al., 1988 Pennisetum americanum (n-2) II + 1V
II Two chromosome additions
1. dTri Frost, 1927 Matthiola incana (n-2) II + 2III
2. DaTri Shahla & Tsuchiya, 1986 Hordeum vulgare (n-1) II + 1IV
3. T Blakeslee, 1921 Datura stramonium (n-2) II + 1IV
4. MteT Gill, 1974 Lycopersicon esculentum (n-2) II + 1VI
5. DteT Gill, 1974 Lycopersicon esculentum (n + 1) II
6. DtT Fedak & Helgason, 1970 Hordeum vulgare (n + 1) II
7. ItT Dhar, unpublished Plantago lagopus (n-1) II + 1IV
8. IDtT Dhar & Koul, 1995 Plantago lagopus (n + 1) II
9. ItTripTri Dhar, unpublished Plantago lagopus (n-2) II + 2III
III. Three or more chromosome additions
1. (trp) Tri Lesley, 1928 Lycopersicon esculentum (n-3) II + 3III
2. (tet) Tri Sai Kumar et al., 1982 Pennisetum americanum (n-2) II + 1III + 1IV
3. IDtTpTri Dhar, unpublished Plantago lagopus (n) II + 1III
4. P Sapre & Barve, 1983 Coix gigantea (n-1) II + 1V
5. ItP Dhar & Koul, 1995 Plantago lagopus (n) II + 1III
6. ItPTri Dhar, unpublished Plantago lagopus (n-1) II + 2III
7. QTri Sapre & Mishra, 1990 Coix lacryma-jobi (n) III
8. H Sapre & Barve, 1983 Coix gigantea (n-1) II + 1VI
9. ItH Dhar & Koul, 1995 Plantago lagopus (n) II + 1IV
10. O Sapre & Naik, 1991 Coix gigantea (n-1) II + 1VIII
B. Compensation aneuploids
Compensating trisomic CTri Khush & Rick, 1967 Lycopersicon esculentum (n-2) II + 1V
1. tiCTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
2. tteCTri Khush, 1973 Triticum spp. (Triticinae) (n-2) II + 1V
3. DiCTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
4. DteCTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-2) II + 1V
5. iteCTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-2) II + 1V
6. dtTripTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 2III
7. dtTriteTri Pantulu & Rao, 1981 Pennisetum americanum (n-3) II + 1III + 1V
8. DtDiT Kamanoi, 1973 Secale cereale (n + 1) II
9. DMtT Pantulu & Rao, 1981 Pennisetum americanum (n-1) II + 1IV
10. DMtTpTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 1III + 1IV
11. DMtTdtTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 1III + 1IV
12. dDtP Pantulu & Rao, 1981 Pennisetum americanum (n-1) II + 1V

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 103
Table 2. (Continued)

Aneuploid Reference Plant species Characteristic meiotic

configuration

C. Deletion aneuploids
1. M Clausen & Goodspeed, 1926 Nicotiana tabacum (n-1) II + I
2. teM Rick, 1943 Petunia spp. (n-2) II + 1III
3. MtD Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II
4. Mt Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + I
5. Dt Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II
6. dMt Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 2I
7. MtMi Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 2I
8. Mi Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + I
9. DtMt Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II + I
10. N Sears, 1941 Triticum spp. (Triticinae) (n-1) II
11. dM Khush, 1973 Triticum spp. (Triticinae) (n-2) II + 2I
12. (trp) M Khush, 1973 Triticum spp. (Triticinae) (n-3) II + 3I
13. dN Sapre & Barve, 1984 Coix gigantea (n-2) II
D. Two-in-one aneuploids
1. MTri Barve & Sapre, 1986 Coix gigantea (n-2) II + I + 1III
2. tiD Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II
3. MiD Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II
4. Di Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II
5. DtTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
6. tiTri Khush, 1973 Triticum spp. (Triticinae) (n-1) II + 1III
7. DiTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n) II + 1I
8. Trit Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
9. Trii Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
10. dMtTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 2III
11. MtTridtTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 2III
12. NTri Matsumura, 1954 Triticum spp. (n-2) II + 1III
13. NT Sears, 1966 Triticum aestivum (n-2) II + 1IV
E. Pseudo-aneuploids
1. dMi Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 2I
2. dtTri Kimber & Sears, 1968 Triticum spp. (Triticinae) (n-1) II + 1III
3. dtTri(I) Pantulu & Rao, 1981 Pennisetum americanum (n-3) II + 1III + 1IV
4. dDt Pantulu & Rao, 1981 Pennisetum americanum (n+1) II
5. ddtTri Pantulu & Rao, 1981 Pennisetum americanum (n-2) II + 2III
6. dDtdtTri Pantulu & Rao, 1981 Pennisetum americanum (n) II + 1III

1. Mono-trisomic MTri: A typical two-in-one aneu-
ploid which is monosomic for one and trisomic
for another chromosome of the complement (2n –
AS.AL + DS.DL).
2. Teloisodisomic tiD: An individual with one chro-
mosome pair missing but having a telocentric and
an isochromosome for one arm of the two missing
chromosomes [2n – 2(AS.AL) + .AS + AS.AS].
3. Mono-isodisomic MiD: An individual deficient for
one chromosome and having an isochromosome
for one arm of the missing chromosome (2n –
AS.AL + AS.AS).
4. Diisosomic Di: A nullisomic having two ho-
mologous isochromosomes for the same arm of
the missing chromosome pair [2n – 2(DS.DL) +
2(DL.DL)].
5. Ditelotrisomic DtTri: A monosomic having a pair
of homologous telocentric chromosomes for one
arm of the missing chromosome [2n – DS.DL +
2(.DL)].
6. Teloisotrisomic tiTri: An individual deficient for
one chromosome and carrying instead a telocen-
tric and an isochromosome for the same arm of
the missing chromosome (2n – DS.DL + DL.DL +
.DL).

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104

Table 3. Proposed new names of some of the aneuploids

Previous name Chromosomal formula Proposed name

1. Double telotrisomic 2n-AS.AL + .AS + .AL Mono double telo

plus primary trisomic + DS.DL disomic plus primary
dtTripTri trisomic MdtDpTri
2. Double telotrisomic 2n-AS.AL + .AS + .AL Mono double telodisomic
plus tertiary trisomic + BS.DL plus tertiary
dtTriteTri trisomic MdtDteTri
3. Dimonotelo tetrasomic 2n-AS.AL +2(.AS) + .AL Mono ditelo telo-
DMtT trisomic MDttTri
4. Dimonotelo tetrasomic 2n-AS.AL-BS.BL + 2(.AS) Nulli ditelo telo
plus double telo- + .AL + .BS +.BL trisomic plus
trisomic DMtTdtTri double telodisomic
NDttTridtD
5. Double ditelopenta- 2n-AS.AL + 2(.AS) + 2(.AL) Mono double ditelo-
somic dDtP trisomic MdDtTri
6. Monotelodisomic MtD 2n-.AS Monotelosomic Mt
7. Monotelosomic Mt 2n- 2(AS.AL) + .AS Nullitelosomic Nt
8. Ditelosomic Dt 2n-2(AS.AL) + 2(.AS) Nulli ditelosomic NDt
9. Monoisosomic Mi 2n-2(AS.AL) + AS.AS Nulli isosomic Ni
10. Monotelo monoiso- 2n-2(AS.AL) + AS.AS + .AL Nulli teloisosomic
somic MtMi Nti
11. Double monotelo- 2n-2(AS.AL) + .AS + .AL Nulli double telo
somic dMt somic Ndt
12. Ditelo monotelosomic DtMt 2n-2(AS.AL) + 2(.AS) + .AL Nulli tritelosomic NTrit
13. Diisosomic Di 2n-2(AS.AL) + 2(AS.AS) Nulli diisosomic NDi
14. Telo-isodisomic tiD 2n-2(AS.AL) + .AS + AS.AS Nulli telo-iso disomic NtiD
15. Ditelotrisomic DtTri 2n-DS.DL + 2(.DL) Mono ditelodisomic MDtD
16. Diisotrisomic DiTri 2n-DS.DL + 2(DL.DL) Mono diiso disomic MDiD
17. Teloisotrisomic tiTri 2n-AS.AL + .AS + AS.AS Mono telo isodisomic MtiD
18. Tritelosomic Trit 2n-2(DS.DL) + 3(.DL) Nulli tritelodisomic NTritD
19. Triisosomic Trii 2n-2(DS.DL) + 3(DL.DL) Nulli triisodisomic NTriiD
20. Double monotelo 2n-AS.AL-BS.BL + 2(.AL) Double mono double
trisomic dMtTri + 2(.BL) ditelo disomic dMdDtD
21. Monotelotrisomic plus 2n-AS.AL-BS.BL + 2(.AL) Double mono tetra
double telotrisomic MtTridtTri + .BS + .BL telo trisomic dM(tet)tTri
22. Double monoisosomic 2n-2(DS.DL) + DS.DS Nulli double iso
dMi + DL.DL disomic NdiD
23. Double telotrisomic 2n-AS.AL + .AS + .AL Mono double telo
dtTri disomic MdtD
24. Double telotrisomic 2n(TBL.DL)-AS.AL Mono double telodisomic
plus interchange dtTri(I) + .AS + .AL plus interchange MdtD(I)
25. Double ditelosomic dDt 2n-2(AS.AL) + 2(.AS) Nulli double ditelo
+ 2 (.AL) disomic NdDtD
26. Double double telo- 2n-AS.AL-BS.BL + .AS + Double mono tetra
trisomic ddtTri + .AL + .BS + .BL telodisomic dM(tet)tD
27. Double ditelosomic plus 2n-2(AS.AL)-DS.DL + Nulli mono double
double telotrisomic dDtdtTri 2(.AS) + 2(.AL) + .DS + .DL ditelodisomic NMdDtD

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7. Diisotrisomic DiTri: A monosomic having in place
of the missing chromosome a pair of homolo-
gous isochromosomes for one arm of the missing
chromosome [2n – DS.DL + 2(DL.DL)].
8. Tritelosomic Trit: An individual missing one chro-
mosome pair but having three homologous telo-
centric chromosomes for one arm of the missing
pair [2n – 2(DS.DL) + 3(.DL)].
9. Triisosomic Trii: An individual deficient for a
chromosome pair and having instead three ho-
mologous isochromosomes for one arm of the
two missing chromosomes [2n – 2(DS.DL) + 3
(DL.DL)].
10. Double monotelotrisomic dMtTri: An individual
missing two non-homologous chromosomes but
having instead four telocentrics; two for an arm of
one chromosome and two for an arm of the other
missing chromosome [2n – AS.AL – BS.BL +
2(.AL) + 2(.BL)].
11. Monotelotrisomic plus double telotrisomic Mt-
Tri dtTri: An individual deficient in two different
chromosomes, but having four telocentrics; two
of which represent an arm of one missing chro-
mosome and one each representing the two arms
of another missing chromosome [2n – AS.AL –
BS.BL + 2(.AL) + .BS + .BL)].
12. Nulli-trisomic NTri: Aneuploids trisomic for one
chromosome and nullisomic for another [2n –
2(DS.DL) + AS.AL].
13. Nulli-tetrasomic NT: Aneuploids nullisomic for
one chromosome and tetrasomic for another [2n –
2(DS.DL) + 2(AS.AL)].
E. Pseudo-aneuploids
This category includes chromosome variants which
do not represent true aneuploids. They may not have
altered chromosome numbers. Even when their chro-
mosome number is altered the gene content is the same
as in disomics. They have been named substitution
diploids by Sybenga (1975). To this category belong
the following types of aneuploids.
1. Double monoisosomic dMi: An individual de-
ficient for a chromosome pair and having in-
stead two isochromosomes, one for each arm of
the missing chromosome pair [2n – 2(DS.DL) +
DS.DS + DL.DL].
2. Double telotrisomic dtTri: A monosomic with
the missing chromosome represented by two telo-
centrics, one for each arm of the missing chromo-
some (2n – AS.AL + .AS + .AL).
105
3. Double telotrisomic plus interchange dtTri(I): A
double telotrisomic in which two other chromo-
somes are involved in a reciprocal translocation
[2n(TBL.DL) – AS.AL + .AS + .AL].
4. Double ditelosomic dDt: An individual deficient
for a pair of homologous chromosomes but having
instead four telocentric chromosomes one for each
missing arm [2n – 2(AS.AL) + 2(.AS) + 2(.AL)].
5. Double double telotrisomic ddtTri: A double
monosomic in which each missing arm is repre-
sented by a telocentric chromosome [2n – AS.AL
– BS.BL + .AS + .AL + .BS + .BL].
6. Double ditelosomic plus double telotrisomic dDt
dtTri: An individual nullisomic for one chromo-
some, monosomic for another and having instead
six telocentrics, each telocentric representing a
missing arm [2n – 2(AS.AL) – DS.DL + 2(.AS) +
2(.AL) + .DS + .DL].
All the types of aneuploids can be identified during
meiosis. Table 2 lists the aneuploids, the plants in
which they were first reported and the characteristic
meiotic configurations by which they can be identified.
In categorizing the aneuploid types emphasis has
been laid on changes in the gene content because of
aneuploidy. Some terms listed above, which have been
coined by earlier workers, on the basis of changes in
chromosome number, have become ambiguous and do
not convey the precise nature of the aneuploid. Such
names have been substituted by new names (see Ta-
ble 3). The first word of these names designates the
magnitude of deletion, the second word refers to the
substitution and the third to the overall status of the
individual.
Acknowledgements
The authors thank the Head, Department of Bio-
sciences, University of Jammu for providing neces-
sary facilities. We are grateful to Dr. J.L. Karihaloo,
NBPGR, IARI New Delhi, for making valuable sug-
gestions. We are also thankful to the reviewers for crit-
ical evaluation and suggestions for the improvement of
the paper.
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