J. Phycol.
24, 203-208 (1988)
DAYLENGTH AND LIGHT RESPONSES IN GROWTH AND FERTILITY OF
GLOSSOPHORA KUNTHII (PHAEOPHYTA, DICTYOTALES) FROM
PACIFIC SOUTH AMERICA’
Alicia J. Ho$mann
Departmento de Biologia Ambiental y d e Poblaciones, Facultad d e Ciencias Biolbgicas, Pontificia Universidad
Cat6lica de Chile, Casilla 114-D, Santiago, Chile
ABSTRACT
Excised ligulae of Glossophora kunthii (C. Ag.) J.
‘4g. were cultured i n photoperiods of 4-24 h and photon
fluence rates of 10-75 pmol. n - 2 . s - 1 . Daylength znter-
acted with zrradiance on the growth of the ligulae. Max-
imal growth of primary ligulae occurred i n long-day reg-
imens with high irradiances suggesting a n effect of
irradiance on photosynthesis and growth. I n contrast,
growth of secondarj ligulae was greatest i n short-daj re-
gimes. Differences uere signiJicant at the hzghest irradi-
ance tested. Diferentiation o f tetrasporangia on the lig-
ulae is a short-day photoperiodic response. Dajlengths of
8.5 h or less induced a sharp increase i n numbers offertzle
ligulae and tetrasporangia attaining maturity Interrup-
tions o f the dark perzod decreased the development of te-
trasporangza; the number of interruptions had a cumu-
lative inhibitory effect. Differentiation of reproductive
structures u1as injuenced bg interactions of photoperiod
and irradiance. Maximum numbers of tetrasporangaa were
formed at short-dug regimes and low irradiances; differ-
entiation ulas completelj inhibited at long-day conditions
and high irradiance.
Kej index words: brown algae; fertility; Glossophora
kunthii; growth; irradiance; night-break; photoperiodic
responses
Glossophora kunthii (C. Ag.) J. Ag. is a brown alga
of subantarctic distribution (Levring 1941, 1960,
Lindauer et al. 1961) that grows in the subtidal and
in tidal pools. The species has an isomorphic life
history, and individual plants are perennial (Sante-
lices and Vera 1984). Seasonal differences observed
in vegetative growth and spore production (Hoff-
mann, unpubl.) suggest that these stages are regu-
lated by environmental conditions. Photomorpho-
genetic responses have been described in relatively
few Phaeophyta (Dring 1984). They include change
from crustose to upright growth and onset of re-
production (Nakahara and Nakamura 197 1 , Dring
and Luning 1975, Bird and McLachlan 19’76,Terry
and Moss 1980, ten Hoopen et al. 1983). Growth
and gametogenesis respond to different irradiances
and to red and blue light interactions in some species
(Dring and Luning 19’75, Muller and Clauss 19’76,
Liining 198 1 , Hoffmann and Santelices 1982, Dring
et al. 1985, Stroemgren 1985). Photoperiod and light
AccPptrd: 1 Februarj 1988.
203
components mainly have been found to control the
development of reproductive structures, although
environmental control of some phases of vegetative
growth also has been described. Moreover, photo-
periodic responses may depend on the level of ir-
radiance (Dring 1984). T h e aim of the present study
was to investigate the role of daylength in fertility
and assess how the interactions of daylength and
irradiance affect vegetative growth and reproduc-
tive structures in Glossophora kunthii.
MATERIALS AND METHODS
Glossophora kunthii forms membranaceous thalli with a few di-
chotomous branches. Plants grow from a large apical cell (Fig.
la, b) to form thalli up to 25 cm long, 6-14 mm wide, and 150-
300 pm thick. These thalli consist of a central layer of large cells,
an epidermal layer and, occasionally, an intermediate layer of
one to three cells (Fig. lc). T h e frond surface bears a large num-
ber of small superficial ligulae that are differentiated near the
thallus apex and grow up to 3-4 mm long. Ligulae grow from
an apical cell, and their morphology is similar to that of the frond
(Fig. Id-f). In older ligulae, at some distance from the thallus
apex, a few epidermal cells at the borders of the ligulae may
differentiate into apical cells and give rise to lateral outgrowths.
Tetrasporangia develop mostly on the surface of the ligulae fac-
ing the frond (Fig. lg) and less frequently on the thallus surface.
Excised ligulae are easily cultured in large numbers.
Mature Glossophora kunthii plants were collected subtidally at
Las Cruces, central Chile (33”15’ S, 71”38’ W) and transported
to the laboratory in plastic buckets filled with seawater. Fronds
were then washed with filtered seawater. Ligulae were excised
with a pair of forceps and examined under a dissecting micro-
scope. Sterile ligulae with only one apical cell (Fig. 2) were se-
lected and incubated in 60 mm Petri dishes containing enriched
seawater (SWM3, McLachlan 1973). Two mL of a stock solution
of 250 mg. L-’ germanium dioxide were added per liter of culture
medium to inhibit diatom growth (Lewin 1966). T h e culture
medium was changed each week without further addition of ger-
manium dioxide, although no damage was detected on the ligulae.
Ligulae were examined weekly to check for epiphyte growth.
Whenever possible, epiphytes were eliminated and heavily epi-
phytized ligulae were discarded.
Culture chambers equipped with cool white fluorescent tubes
(Philips 40W) and set at 8:16, 12:12 or 16:8 h LD cycles were
used for the experiments. Different irradiances were obtained by
varying the distances to the light sources; photon fluence rates
were measured using a LI-COR Model 185-A quantum meter.
Temperature was set at 15” C ( ? l o C) and monitored by ther-
mometers with their bulbs immersed in 100 mL distilled water.
T h e interactive effects of daylength and it-radiance were tested
at 8:16, 12:12 or 16:8 h LD with photon fluence rates of 10, 25,
50, or 75 fimol.m-2.s-1.Two replicate dishes were used for each
condition tested. Results were evaluated on day 30 of the exper-
iments. After eliminating all ligulae on which epiphytes had de-
veloped, the following parameters were measured on 15 ligulae
per dish in each experimental condition: (i) number of new apical
204 ALICIA J. HOFFMANN
-
-1
I similar in the three photoperiods. After one week,
FIG.1. Morphological structure of Glossophora kunthii. (a) frond,
(b) apical cell, (c) frond cross-section with central cell layer and
epidermal cell layer, (d) ligula bearing tetrasporangia, (e) apical
cell of ligula, (f) cross-section of ligula, (g) epidermal layer in
front view, (h) tetrasporangial initial, (i) tetrasporangial initial
before division, (j) tetraspores, (k) released spores. Scale bar for
g-k = 100 pm.
cells differentiating on the margin of the ligulae; (ii) area of the
ligulae; (iii) number of ligulae bearing tetrasporangia, numbers
of tetrasporangia per ligula and proportions of tetrasporangia at
different stages of maturity. The following stages were distin-
guished: (stage 1 ) tetrasporangial initials werejust discernible due
to their larger size, as compared to epidermal cells; (stage 2)
tetrasporangial initials had turned dark and reached their max-
imum size before dividing; (stage 3) cruciate division had taken
place. The area of the ligulae was calculated with a polar com-
pensating planimeter (Los Angeles Scientific Instrument C o . , Los
Angeles) on contours obtained with a camera lucida. Primary and
secondary ligulae areas were measured separately.
T o determine the critical daylength for fertility response, pho-
toperiods of 4 , 6 , 8 , 10, 14, 18 and 20 h daylength were obtained
by placing light-tight boxes over the culture dishes during dark-
ness and removing them at the hours when dishes had to be
exposed to light. One set of dishes was exposed to continuous
light. The culture dishes received a photon fluence rate of 10
pmol.m-2.s-1. Four culture dishes containing 30 ligulae each
were used as replicates.
To test for short-day responses, the dark period in an 8: 16 h
LD cycle was interrupted in the middle by one-hour white light
periods. The light had the same photon fluence rate (10 pmol.
m-2. s-I ) as the main photoperiod. In order to test for cumulative
effects of light interruptions, four treatments with different num-
bers of interruptions of darkness were set. In the first treatment
the dark period was interrupted on the first night only; in the
second, all periods of darkness were interrupted during one week;
in a third set, dark periods were interrupted during two weeks;
and, in the fourth, dark periods were interrupted throughout
the three weeks of the experiment. Dark periods were not in-
terrupted in the control treatment.
Results were analyzed using factorial analysis of variance (AN-
OVA) after angular transformation whenever pertinent. Inter-
actions were tested using Tukey’s test (Sokal and Rohlf 1981);
only the significant differences (P < 0.05 for both tests) are dis-
cussed in the text.
RESULTS
Plants were either tetrasporophytic or sterile.
Much higher percentages of tetrasporophytes have
been recorded in nature for several species of the
Dictyotales (Wynne and Loiseaux 1976).
After about one week in culture, some epidermal
cells of the ligulae differentiated into apical cells
(Fig. 3). T h e number of new apical cells increased
under higher irradiances in the three photoperiods
tested (Fig. 6). ANOVA showed a significant inter-
action of the effects of photoperiod and irradiance
on the number of apical cells that developed. T h e
largest numbers of new apical cells differentiated in
the 12: 12 h LD cycle under all irradiances. Tukey’s
test showed a significant effect of irradiance on the
numbers of cells at 10 and 25 pmol.rn-*.s-l. At 75
pmol-m-2.s-1 the number of new apical cells was
the apical cells began to divide and gave rise to sec-
ondary ligulae (Fig. 4).
Although the primary ligulae showed some growth
under all culture conditions (Fig. 7A), their area
depended on photoperiod, irradiance, and their in-
teractions (ANOVA). T h e largest primary ligulae
developed in the 12:12 h LD cycle under all irra-
diances tested, although no significant differences
were found with ligulae incubated in a long-day pho-
toperiod. T h e ligulae grew significantly less in the
short-day photoperiod under irradiances between
10 and 50 pmol.m-2.s-1 (Tukey’s test). In contrast,
at 75 pmol.m-2.s-1 ligulae attained the same size as
in the other photoperiods.
T h e area of the secondary ligulae was also affected
by photoregime (Fig. 7B). In the three photoperiods
tested, the area tended to be larger under increasing
irradiances. Under 75 pmol.m-2.s-1, ligulae cul-
tured in the short-day photoperiod grew signifi-
cantly more than those cultured in the 12:12 and
16:8 h LD cycles (Tukey’s test).
Although after four weeks in culture both pri-
mary and secondary ligulae had grown more under
the highest irradiance than under the other irradi-
ances tested, most of the ligulae had turned pale as
compared to those grown under lower irradiances.
Tetrasporangia began to differentiate on the lig-
ulae after approximately two weeks in culture (Fig.
4). Some of the epidermal cells underwent a peri-
clinal division. T h e distal cell (tetrasporangial initial)
darkened, increased in volume and projected out
from the epidermis (Fig. 5). Cruciate tetrasporangia
 DAYLENGTH A N D LIGHT O N G. KUNTHII 205

0.5mm

FIGS.2-5. Developmental stages of ligulae. FIG. 2. Isolated ligula, showing large apical cell (arrow). FIG. 3. Cross-section of a ligula
showing central layer of large cells and epidermal layer. One epidermal cell (arrow) is probably changing into a secondary apical cell.
FIG. 4. Ligula with lateral outgrowths (“secondary” ligulae) that develop after approximately three weeks in culture. Tetrasporangia
are differentiating on the epidermal surface. FIG. 5. Tetrasporangium developing on the epidermal layer. Stalk cell is visible.

developed (Fig. 1 h-j). Finally the cells separated,
and four spores were released (Fig. 1k).
T h e development of reproductive structures was
influenced by photoperiod. At daylengths between
4 and 8.5 h, tetrasporangia differentiated in SO-SO%
of the ligulae. In contrast, daylengths over 8.5 h
induced a sharp decline in fertility, with about 15%
fertile ligulae (Fig. 8). Results show that the critical
daylength is about 8.5 h for the fertility response
and that longer daylengths reduce the differentia-
tion of tetrasporangia. In addition, the degree of
maturity of the reproductive structures varied with
daylength. ANOVA showed a significant effect of
photoperiod on the stage of maturity reached by
tetrasporangia. At 4-8 h daylengths, 60-65% of the
reproductive structures attained stage 2 or 3, and
some spores were released after 28 days in culture
(Fig. 9). Tukey’s test showed a significant effect of
4,6 or 8 h daylength on the percentages of mature
tetrasporangia. In 12 h daylength, around 25% of
the tetrasporangia attained stage 2 or 3. Between
14 h and 18 h, most reproductive structures re-
mained at stage 1, and very few tetrasporangia
reached maturity.
Daylength also influenced the number of tetra-
A - PRIMARY LIGULAE

:
-I

3
t, I I I

-
0
A
8 - SECONDARY LIGULAE
Y 41

1 I I
I 1 I 10 25 50 75
u)
Z 10 25 50 75

IRRADIANCE ( p mol m-2 s1 )

FIG. 6. Interactions of daylength and irradiance on differ-
entiation of new apical cells on ligulae of G. kunthii after 30 days
in culture. 16:8 h LD cycles (0);12:12 h LD cycles (0);8:16 h
LD cycles (A). Each point represents the mean for 30 measure-
rnents.
FIG. 7A, B. Interactions of daylength and irradiance on the
growth of (A) “primary” and (B) “secondary” ligulae of G. kunthzz
after 30 days in culture. 16:8 h LD cycles (0);12:12 h LD cycles
(0);8: 16 h LD cycles (A). Each point represents the mean for 30
measurements.
206 ALICIA J. HOFFMANN

PHOTOPER100 TETRASPORANGIAL DIFFERENTIATION I X ,

EP7

801

57 0

4b 2

49

1ETRASPORANGIA

8 16 87i301130

FIG. 10. Effects on differentiation of tetrasporangia of one-

hour interruptions in the middle of the dark period in a 8:16 h
LD cycle. Interruptions were performed the first night only, or
during one to three weeks throughout the experiment. The up-
I I 77-
permost line represents the control treatment, with uninter-
rupted dark periods.
4 6 I
8
I
Ib 12 f4
1 I
16 18 20 24
DAYLENGTH ( h )
One-hour light interruptions of the dark period
FIG.8. Effects of daylength on tetrasporangial differentiation.
Each value represents counts of about 140 ligulae. Vertical bars
significantly decreased the differentiation of tetra-
are 95% confidence limits. sporangia in ligulae incubated under a short-day
photoperiod (ANOVA). However, the magnitude of
the inhibition depended on the number of inter-
sporangia reaching maturity. After 30 days in cul- ruptions (Fig. 10). In the control experiment, where
ture, under 8 h daylength the average number of the dark period was not interrupted, tetrasporangia
mature tetrasporangia per ligula was 5.52 f 0.66 developed on almost 90% of the ligulae after 30 days
( f S D ) , under 12 h daylength, 0.39 k 0.41, and un- in culture. When only the first dark period was in-
der 16 h daylength, 0.83 f 0.29. terrupted, 80% of the ligulae became fertile. When
the dark periods were interrupted in the course of
one or two weeks, fertility dropped to 60% and 46%,
respectively; and when the dark periods were inter-
rupted during three weeks, only 5% of the ligulae
became fertile.
T h e differentiation of reproductive structures was
lo]80

4 6 8 ?O 12 14 16 18
DAYLENGTH ( h )
10 25 50 75

I RRAD I ANCE ( p m o l mi’<’

FIG.9. Effects of daylength on stage of development attained FIG. 11. Interactions of photoperiod and irradiance on at-
by tetrasporangia. Each bar represents % tetrasporangia attaining tainment of fertility (differentiation of tetrasporangia) of G. kun-
stages 1, 2, or 3 under different daylengths, after 30 days in thii after 30 days in culture. 16:8 h LD cycles (0);12:12 h LD
culture. cycles (0);8: 16 h LD cycles (A).
 DAYLENGTH AND LIGHT O N G.KUNTHII
10 25 50 75 10
I I 1
I I
16:8h LD
FIG. 12. Interactions of daylength and irradiance on growth of “primary” and “secondary” ligulae and on fertility (differentiation
of tetrasporangia) of G. kunthii. Each circle represents 10% of fertile ligulae.
significantly influenced (ANOVA) by the interac-
tions of photoperiod and irradiance (Fig. 11). In the
8:16 h LD cycle a higher proportion of the ligulae
became fertile when compared to the other pho-
toperiods under all irradiances tested. Fertility de-
pended on irradiance (Tukey’s test): the highest per-
centage of fertility occurred under low irradiance
(10 pmol .mP2.s-I).
In the 12:12 h LD cycle, tetrasporangia differ-
entiated on 20-30o/c of the ligulae, depending on
irradiance, but no clear effects were recognizable
on fertility under different amounts of irradiance.
In the 16:8 h LD cycle, very few ligulae became
fertile, some tetrasporangia developed but their
maturation depended on irradiance. Under 10 pmol.
m-2.s-1, 8 ligulae (14.6%) became fertile. Tetra-
sporangial development began in 36 cells of the 8
ligulae, but only 22% reached stage 3. Under 75
pmol .m-2.s-1, differentiation of tetrasporangia be-
gan in 22 cells of 7 ligulae (9.5%), but the tetraspo-
rangial initials remained pale and none developed
beyond stage 1.
DISCUSSION
Glossophora kunthii has a well defined critical day-
length in the formation of tetrasporangia. This re-
sponse is inhibited by short light-breaks in the mid-
dle of dark periods. At least 15.5 h darkness are
required in order for over 50% of the ligulae to
become fertile. Most of the responses reported in
brown algae are short-day mediated (Dring 1984).
Interruptions of the dark period had a cumulative
effect: the interruption of one dark period resulted
in a 10% decrease in fertility, whereas further in-
terruptions progressively decreased the proportions
of fertile ligulae.
Daylength and irradiance had varying effects on
growth and fertility, as shown in Figure 12. New
207
25 50 75 10 25 50 75
( p mol m-2 s- 1 )
I
12:12h L D 8:16h LD
apical cells differentiated under all conditions tested,
possibly reflecting a release from the dominance ex-
erted by the apical cell of the thallus from which
ligulae were obtained. T h e existence of such apical
dominance was demonstrated in several algal species,
among them some Dictyotales (Russell 1970, Dahl
1971, Moss 1974, Ducreux 1977, Gaillard and
L’Hardy Halos 1980). Primary ligulae grew more
under long-day and neutral regimes with high ir-
radiance than under short-day regimes with low ir-
radiance. This is probably due to the effect of day-
length and irradiance on photosynthesis and growth
rather than to a photoperiodic response. In contrast,
the secondary ligulae grew significantly more in
short-day than in long-day regimes, suggesting that
a photoperiodic response could be involved in this
stage of development.
Apparently high irradiance not only influenced
the growth of ligulae but also some specific devel-
opmental processes, resulting in decoloration of the
ligulae. In Dictjota dichotoina (Hudson) Lamouroux,
irradiation with wavelengths over 500 nm resulted
in fading of the thalli accompanied by a dark col-
oration of the tip region (Miiller and Clauss 1976).
Tetrasporangia began to differentiate under most
experimental conditions, but the number and de-
gree of maturity reached were regulated by the in-
teractions of photoperiod and irradiance. A higher
number of tetrasporangia differentiated under short-
day regimes and low irradiance than under the other
conditions tested. Long-day photoperiods, or high
irradiance, apparently hindered the attainment of
full maturity.
Photoperiodic regulation of development has been
demonstrated in about 40 algae, among them nine
Phaeophyta (Dring 1984). So far, all brown algae
with photoperiodic responses have heteromorphic
life histories (Nakahara and Nakamura 1971, Dring
208 ALICIA J. HOFFMANN
and Luning 1975, Luning 1982, Lobban et al. 1985).
Glossophora kunthii seems to be the first perennial
brown alga with an isomorphic life history to show
photoperiodic regulation of development. In Rho-
dophyta, photoperiodic control of development also
has been shown in some species with isomorphic life
histories, e.g. Dumontia contorta (S. G. Gmelin) Ru-
precht, Hal)lmenia latqolia Kutzing, and Gigartina aci-
cularis (Roth) Lamouroux (Rietema 1982, Rietema
and Breeman 1982, Guiry 1984, Guiry and Cun-
ningham 1984). In most algae, photoperiodic re-
sponses are restricted to the sporophyte generation;
up to now, the red algae Gigartina acicularis and
Helminthora divaricata J. Agardh are the only species
where photoperiodic control has been reported for
both generations (Guiry 1984, Guiry and Cunning-
ham 1984, Cunningham and Guiry 1985). Further
studies on G. kunthii are needed to determine wheth-
er both generations are under photoperiodic con-
trol.
The author wishes to thank S. Contreras, C. Orellana, G. Mufioz
and A. Morgado for field collection of subtidal material. M. E.
Malbrin participated as research assistant. L. Gonzilez is thanked
for technical assistance. Dr. B. Santelices' critical reading of the
manuscript and the valuable comments of two anonymous re-
viewers are greatly appreciated. This work was supported by
grants DIUC 97/87 (P. Universidad Cat6lica de Chile) and FON-
DECYT 7 18/87.
Bird, N. L. & McLachlan, J. 1976. Control of formation of re-
ceptacles in Fucus distichus L. subsp. distichus (Phaeophyceae.
Fucales). Phycologia 15:79-84.
Cunningham, E. M. & Guiry, M. D. 1985. Photoperiodic and
temperature control in the life history of Helminthora divari-
cata (Rhodophyta) from Ireland. Int. Phjcol. Congress (Abstr.)
2:34.
Dahl, A. L. 1971. Development, form and environment in the
brown alga Zonaria farlowii (Dictyotales). Bat. Mar. 14:76-
112.
Dring, M. J. 1984. Photoperiodism and phycology. Progr. Phycol.
RCS.3~159-92.
Dring, M. J. & Luning, K. 1975. A photoperiodic response me-
diated by blue light in the brown alga Scytosiphon lomentaria.
Planta (Berl.) 125:25-32.
Dring, M. J., Maggs, C. A. & Butler, D. M. 1985. Long-term
growth responses of brown algae to blue and red light. Int.
Phjcol. Congress (Abstr.) 2:39.
Ducreux, P. 1977. Etude experimentale des correlations et des
possibilitks de regheration au niveau de l'apex de Sphace-
laria cirrhosa Ag. Ann. Sci. 'Vat. Bot. 18:163-84.
Gaillard, J. & L'Hardy-Halos, M-T. 1980. Croissance et rami-
fication chez le Dictyota dichotoma (Huds.) Lamouroux (Phko-
phycke, Dictyotales). Phycologza 19:159-67.
Guiry, M. D. 1984. Photoperiodic and temperature responses
in the growth and tetrasporogenesis of Gigartina acicularis
(Rhodophyta) from Ireland. Helgol. Meeresunters. 3 8:3 35-47.
Guiry, M. D. & Cunningham, E. M. 1984. Photoperiodic and
temperature responses in the reproduction of north eastern
Atlantic Gigartina acicularis (Rhodophyta: Gigartinales).
Phjcologia 23:357-67.
Hoffmann, A. J. & Santelices, B. 1982. Effects of light intensity
and nutrients on gametophytes and gametogenesis of Lesso-
nia nigrescens Bory (Phaeophyta). J . Exp. Mar. Biol. Ecol. 60:
77-89.
Levring, T . 1941. Die Meeresalgen der Juan Fernandez Inseln.
In Skottsberg, C. [Ed.] The Natural History of Juan Fernandez
and Easter Island. Almquist & Wiksells Boktyyckeri. Uppsala,
pp. 601-70.
- 1960. Contribution to the marine algal flora of Chile.
Lunds Universitets Arsskrft N.F. Avd. 2 Bd 56 N. 10, C. W.
K. Gleerup, Lund, 83 pp.
Lewin, J. 1966. Silicon metabolism in diatom growth. V. Ger-
manium dioxide, a specific inhibitor of diatom growth. Phyco-
logia 6:l-12.
Lindauer, V. W., Chapman, V. J. & Aiken, M. 1961. T h e marine
algae of New Zealand. 11. Phaeophyceae. Nuzla Hedwigia 3:
129-350.
Lobban, C. S., Harrison, P. J. & Duncan, M. J. 1985. The Phys-
iological Ecolog)' of Seaweeds. Cambridge University Press, Lon-
don, 242 pp.
Luning, K. 1981. Light. In Lobban, C. S. & Wynne M. [Eds.]
The Biologj of Seaweeds. Botanical Monographs 17:326-54.
1982. Photoperiodisch kontrollierte Bildung des neuen
Phylloids bei der Braunalge Laminaria hyperborea. Jahr. Ber.
Bid. Anstalt Helgola?ad 1981:28-9.
McLachlan, J. 1973. Growth media-marine. In Stein, J. R. [Ed.]
Handbook of Phycologzcal Methods. Cambridge University Press,
Cambridge, pp. 25-52.
Moss, B. L. 1974. Morphogenesis. In Steward, W. D. P. [Ed.]
Algal Physiolug)' and Biochemistry. Blackwell, Oxford, pp. 788-
813.
Muller, S. & Clauss, H. 1976. Aspects of photomorphogenesis
in the brown alga Dictjota dichotoma. 2. Pflanzenphysiol. 78:
461-5.
Nakahara, H. & Nakamura, Y. 1971. T h e life history of Des-
inarestia tabacoides Okamura. Bot. Mag. (Tokyo) 84:69-75.
Rietema, H. 1982. Effects of photoperiod and temperature on
macrothallus initiation in Durnuntia contorta (Rhodophyta).
Mar. Ecol. Prog. Ser. 8:187-96.
Rietema, H. & Breeman, A. M. 1982. The regulation of the life
history of Dumontia coiztorta in comparison to that of several
other Dumontiaceae (Rhodophyta). Bat. Mar. 25:569-76.
Russell, G. 1970. Rhizoid production in excised Dictyota dichot-
oma. Brit. Phjcol. J. 5:243-5.
Santelices, B. & Vera, M. E. 1984. Variacibn estacional de floras
marinas en Caleta Horcbn, Chile central. Phycologia Latino
Americana 2533-101.
Sokal, R. R. & Rohlf, F. J. 1981. Biometry W. H. Freeman & Co.,
San Francisco, 859 pp.
Stroemgren, T . 1985. Effects of light on apical length growth
of Fucus distichus ssp. edentatus (Phaeophyta). Mar. Bid. (Berl.)
86:263-70.
ten Hoopen, A,, Bos, S. & Breeman, A. M. 1983. Photoperiodic
response in the formation of gametangia of the long-day
plant Sphacelaria rigidula (Phaeophyceae). Mar. Ecol. Prog.
Ser. 13:285-9.
Terry, L. A. & Moss, B. L. 1980. T h e effect of photoperiod on
receptacle initiation in Ascophyllum nodosum (L.) Le Jol. Br.
Phjcol. J . 15:291-301.
Wynne, M. J. & Loiseaux, S. 1976. Recent advances in life history
studies of the Phaeophyta. Phycologia 15:435-52.