P a r a m e c i a 1411
When large differences in allele frequency are
found among different subgroups within a population
two conclusions follow:
1. Mating is not random across subgroups. If it
were, allele frequencies would not differ among
them.
2. Alleles are not associated randomly within indi-
viduals. Alleles common within a particular sub-
group are found together more frequently than
expected if alleles were randomly associated across
the entire population.
Thus, humans do not form a panmictic population,
although many subgroups of humans form panmictic
subpopulations. Panmixis is always defined relative to
a particular population. With respect to the entire
species, humans are not panmictic. With respect to
many subgroups of the human population, humans
form panmictic subpopulations. It is also important
to remember that the human species departs far less
from panmixis than populations of many other plants
and animals. The average genetic difference among
subpopulations of humans is much smaller than the
average difference among individuals within sub-
populations.
The most extreme departure from panmixis is
found in some ferns and their relatives. In these plants
a free-living haploid generation produces both sperm
and egg, and in some species sperm and egg produced
by the same haploid individual fuse to form a diploid
zygote that is completely homozygous. Many flower-
ing plants and some snails are only a little less extreme
in their departure from panmixis. Diploid hermaph-
rodites produce both sperm and egg that fuse to
form zygotes heterozygous at only half as many
loci as the individual that produced them, on average.
Even animals and plants with separate sexes may
inbreed to some extent, as when cousins mate, broth-
ers mate with sisters, or aunts mate with nephews.
Whether the departure from panmixis is a result of
inbreeding, as just described, or assortative mating, the
effects are similar: alleles with a similar effect are more
likely to be found together within individuals than
expected.
Disassortative mating is less widely recognized, but
may be almost as common as inbreeding and assort-
ative mating. In many flowering plants, for example,
individuals that share alleles at the self-incompatibility
locus are prevented from mating. Only individuals
that carry different alleles at this locus are able to
mate. Similarly, there is evidence in mice that mating
occurs preferentially among individuals with different
genotypes at major histocompatibility complex loci.
With disassortative mating, alleles with a similar effect
are less likely to be found together within individuals
than expected.
See also: Demes; Hardy±Weinberg Law;
Inbreeding; Wahlund Effect
Paralogy
W Fitch
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.0951
The circumstance in which two homologous genes
diverge following gene duplication so that the com-
mon ancestor of the two sequences predates their
cenancestor.
See also: Cenancestor; Orthology; Xenology
Paramecia
K J Aufderheide
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.0952
Since Tracy Sonneborn demonstrated the existence of
a genetic system in Paramecium in the 1930s, members
of this genus have been valuable research organisms
for many types of Mendelian and non-Mendelian
genetic studies. A few dozen species in this genus
have been found throughout the world, mostly in
freshwater habitats. Cells are very large, ranging in
size from about 100 to 300 mm in length, and about
20 to 50 mm in width. In spite of their size, the cell
cycle time is typically short: as little as 5 h for small
species. Paramecia share a number of common cilio-
phoran traits, including the cortex, the elaborate array
of cytoskeletal and membranous structures organized
around the basal bodies of the several thousand cilia
on the cell surface, and nuclear dimorphism, the pos-
session of two distinct types of nuclei in each cell. The
small micronuclei are diploid, do not appear to be
transcriptionally active, and contribute little to the
phenotype of the cell. During mitosis or meiosis,
the micronuclei show condensation of chromosomes
and formation of a spindle apparatus without dis-
solution of the nuclear envelope. Depending on the
species, a cell might possess 1, 2, or 4 or more micro-
nuclei. The large macronuclei are polycopy (i.e.,
containing about 1000 copies of each gene), are
transcriptionally active, and determine most of the