The flower producing tissue of a branch of Kramerta is therefore more likely
to have been exposed more uniformly in time than a branch with a flower head
in Machaeranthera, and quite possibly with larger doses for a position equidistant from the radiation source for the following reason.
Krameria is more
likely to form its own clumps, independent of nearby species compared to
Machaeranthera, which often grows under, or within, shrubs with heavier woody
stems such as Lyctwn and Ephedra,
This particularly contributes to the
uncertainty of dose

to any particular

tissue within Maehaeranthera shrubs.

We note in passing that differences in the rate of occurrence of chromosome
aberrations at two dose levels for Kramerta is significantly g-eater at higher
doses than at lower doses by conventionally accepted statistics.
There are still other conditions which support the hypothesis that chromosomal
aberrations should be expected even in the low-level radiation areas:
1) High interphase chromosome volumes.
Precisely the characteristics which render a plant suitable for cytological analysis, i.e., a smal] number of large
chromosomes with a high interphase chromosome volume, characterizes the more
radiosensitive species (Sparrow, 1962; Sparrow, et al., 1965). All three
species tested here have relatively large chromosomes and should therefore be
among the more radiosensitive,
Wallace and Romney, 1972 repored that

Machaeranthera tortifolia and Atriplex spinescens were the most sensitive,

based on phenological or morphological changes, of 31 NTS species studied
using gamma radiation in the laboratory. We have already noted that Xrwneria
parvifolia in Rock Valley was one of the shrubs responding to increasing
exposure rates with reduction in shrub size, more dead stem material, and
less foliage on live stems (Vollmer and Bamberg, 1975).

2) Artemisia, particularly, may be more sensitive to radiation effects for yet
another reason.
The meiotic division was found to be the most radiosensitive
stage in the plant's life cycle (Sparrow, 1951); and a pliant which has a long
meiotic period would probably be still more sensitive,
Such is likely the
case for Artemisia which breaks dormancy very early in the spring (it ts among
the first to do so); and because of the coo] conditions the meiotic cycle is
likely to be longer than for other, later-flowering, species.
Thus there
appears to be a longer time than for other shrubs during which Artemisia may
be radiation sensitive.

Unlike carefully controlled laboratory studies of radiation induced chromosome
aberrations, it is not possible to know in which stage of the cell cycle or
in what part of the plant life cycle the actual radiation "hits” occurred.
Sparrow, et al., (1961) noted that in chronic irradiation effects, most observed
damage is a result of the previous one or two cell divisions leading to lethal
conditions produced by fragments and dicentrics. Aberrations such as heterozygous inversions and translocations, however, are not lethal in somatic tissue
because the complete complement of genetic material is maintained, although in
These structural rearrangements sometimes produce morphoa different order,
logical or physiological changes by altering linkage groups (Swanson, et al.,
1967). We noted the occurrence of a high frequency of aberrants indiciating
a heterozygous inversion in Machaeranthera (Table 2, Shrub Sample #68), and a
a similar high frequency indicating a translocation in Artemisia (Table 1,
Shrub Sample #46)
If indeed there are radiation induced chromosomal aberrations in Site D,
Area 11, according to our hypothesis there is some data which allows some
predictions as to long-term ecological effects.
Chromosomal aberrations can
lead to a general decline in productivity and eventual disappearance of radiosensitive species in an ecosystem (Garrett, 1967; Woodwefl and Sparrow, 1965;
Woodwell, 1962; Woodwell and Whittaker, 1968). Wowever, radiation has long
been utilized as a method of producing beneficial mutations in crop plants
(Smith, 1958; William and Scully, 1961); and it is possible that penetic
changes could occur as a result of radiation which might establish individuals
or species better adapted to their environment. These adaptations could
include radioresistance due to polyploidy, smaller chromosomes, asexual
reproduction, and faster mitosis (Sparrow and Evans, 1961; Evans and Sparrow,
1961). With such mutations the resistant species might expand and become
dominant.
Finally, whether one accepts the hypothesis that there are more than normal
chromosome aberrancies in Site D, Area 11, or rejects it depends on interpretation of statistical criteria.
The argument has been presented that large
varlability should be expected on biological and environmental conditions,
and that producing conclusively adequate statistical data requires very large
samples which verge on unfeastbility because of the complexity of cytological
procedures.

3) Environmental stresses are known to intensify radiation damage, stresses
such as heat, cold, and drought (McCormick and Platt, 1962; Woodwell, 1962) to
which the NTS species are all subjected.
A few words should be said about the detectability of chromosome aberrations.
There are no doubt many aberrations that will remain undetected because they
occur in cells whose chromosomes are insufficiently spread out for cytological
analysis.
Some aberrations cannot be detected.
These include pericentric
inversions, homozygous inversions and translocations, heterozygous paracentric
inversions in which crossover did not occur within the inverted loop, and
inversions with two strand double cross-over within the inversion loop. Also
fragments may be hidden in a group of migrating chromosomes, dicentrics may
break early in anaphase, and multivalents are sometimes difficult to distinguish
from superimposed bivalents.
For all these reasons, the percentages of aberrations detected probably do not present more than an uncertain fraction of
the total.
Detectable aberrancies are, however, generally acceptable as
comparative basis for analyzing populations.
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