maintained between 23° to 24°C, Energies of illumination were as high as
8.5

X

10° erg cm-? sec’ in visible

light and 6.9 xX 10° erg cm~® secin
red light. At appropriate time intervals,

samples were removed and plated on

agar as described previously (70). The
number of colonies were counted after
a 7-day incubation at 26°C under 880
jJumen/m?.
Euglena

gracilis

cells

illuminated

with white light under aerobic conditions demonstrate a photosensitized oxi-

(17). Even after a 90-minute exposure

indication that the spontaneous muta-

to this intense laser beam, no cell death

tion rate was less than 0.00003 percent.

could be demonstrated, and MacMillan

et al. (17) concluded that the cells did

not contain a photosensitizer which
could absorb at 632.8 nm. As shown
below, however, a broader source of

red light can bring about photosensitization in the absence of exogenous photosensitizers in E. gracilis.
White mutant colonies developed
from cells surviving either high light
intensities (8.5 X 10% erg cm-? sec-!) or

dation or bleaching of chlorophyll.

intensities equivalent to bright sunlight

ing process in air consists of an initial

No such white colonies were observed
when cells were exposed to the optimum light intensity for chlorophyil

Under anaerobic conditions there is no
loss of pigment (Fig. 1). The bleachlag period followed by a rapid disappearance of chlorophyll. This lag period
can be shortened or abolished by the
use of 95 percent Oz and 5 percent
CO, as the gas phase. These patterns
are similar to those described for
Chlorella pyrenoidosa

(4.5 & 10° erg cm? sec!) (Table 1).

syothesis (3 X 10° erg cm-? sec"'}, an

deficient bacteria, Sistrom et al. (/2)

have suggested that this mechanism involves carotenoid pigments acting as
“chemical buffers” to protect cells
against photosensitized oxidations. In

phenomenon.

These white colonies could be subcultured on either agar plates or in
liquid medium, giving rise to cells
which are identical in appearance to the
albino mutants induced by ultraviolet
light, heat, or streptomycin (/8).
Fluorescence microscopy did not reveal

the presence of any red fluorescence,
indicating not only the absence of chloroplasts containing chlorophyll but also
of proplastids containing protochloro-

Table 1. Mutagenic effect of visible light in Euglena gracilis var. bacillaris grown in air.

White-light
irradiation

(11). The lag

period presumably represents the time
during which the normal protective
mechanism is still capable of operating.
Based on their studies with carotenoid-

High-intensity rad light (> 610 nm)
also led to the development of white
mutant colonies, a strong suggestion
that the endogenous chlorophyll functions as the photosensitizer for this

erg cm-* sec7!

Cells
plated

<
*
*
x

(No.)

28,000

28,000

0

6
2
5
5

60,000
27,000
27,000
60,000

150
2,000
281
127

75
15
20
63

10°
10°
10°
108

*Red light (> 610 nm) used to irradiate sample.

(/4) reduced antheraxanthin to zeaxan-

thin at a rate commensurate with the
epoxidation of zeaxanthin to antheraxanthin by visible light and O2 (J5).

Under conditions of high light intensity,

this protective mechanism could no
longer cope with the rate of oxidation,
and various effects of aerobic photo-

sensitivity, including pigment bleaching

and death would occur.

The duration of the lag period seen

in Fig. 1 and the time needed for complete

bleaching depend on light

tinuous-wave gas laser (29 mw) emitting at 632.8 nm in the absence of
exogenous photosensitizers, have failed

ao AIR

———
a

=e

“a

ao

100

mel

2

ERcS/cm /SEC
meee 8,5 X 10°

80

—-=—=3,0x10°

60 F

407

20

in-

tensity, ceil concentration, and the
growth phase of the culture. Cells in
the early exponential phase bleach more
rapidly than do cells in the late exponential phase. Similar results have
been reported for photosensitization in
Rhodotorula glutinis (16).
Previous attempts to photosensitize
E. gracilis, with the light from a con-

8 DECEMBER 1967

120F

RELATIVE CHLOROPHYLL CONCENTRATION (2)

mechanism would only remain effective

as long as antheraxanthin deepoxidase

(No.)

(No.)

6

E. gracilis, Krinsky (13) has proposed

that the carotenoids zeaxanthin and
antheraxanthin (5,6-epoxy-zeaxanthin)
function as protective pigments, This

White mutant
colonies

Hours

3.0 < 105
8.5
4.5
4.5
*6.9

Colonies
surviving

~
0

x
i

0

1

1

rt

j

2

l

j

3

I

L

4

I

5

i

L

6

HOURS OF FLLUMINATION
Fig. 1. The effect of tlumination with high-intensity visible light on the relative
chlorophyll content of Euglena gracilis var. bacitlaris cells in the logarithmic phase of
growth exposed to a nitrogen atmosphere (MI), air (©), or 95 percent O, and 5 percent
CO, atmosphere (X). The control sample (@) was exposed to 3 x 10° erg cm™ sec*
white light (the optimum light intensity for chlorophyll synthesis) in an atmosphere of
air, whereas the other samples were exposed to 8.5 x 10° erg cmsec”? white light.
At the indicated times, samples were removed and analyzed for chlorophyll as described
by Arnon (28). The results are expressed relative to the concentration of chlorophyll

present at the beginning of the experiment.

q

1333

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