全 文 :doi: 10.7541/2016.72
MORPHOLOGY, GROWTH AND PHOTOSYNTHETIC RESPONSES OF THE
CYANOBACTERIUM ARTHROSPIRA PLATENSIS TO DIFFERENT
WAVEBANDS IN SOLAR SPECTRUM
MA Zeng-Ling1, M. Arocena Joselito1, 2 and GAO Kun-Shan3
(1. Zhejiang Provincial Key Laboratory for Subtropical Water Environment and Marine Biological Resources Protection, Wenzhou
University, Wenzhou 325035, China; 2. Environmental Science and Engineering, University of Northern British Columbia, Prince
George V2N4Z9, Canada; 3. State Key Laboratory of Marine Environmental Science, Xiamen University, Xiamen 361005, China)
Abstract: Light is known to regulate morphological development and photosynthetic performance of the eco-
nomically important cyanobacterium, Arthrospira platensis. However, light quality under different
wavelengths on its growth and physiology is yet to be understood. In this study, we grew A. platensis D-0083
trichomes in quartz tubes covered with different cutoff foils and one band-pass filter, so that the cells re-
ceived different wavebands of irradiances, and measured its growth, morphology and photosynthetic perfor-
mances. Spirals of A. platensis D-0083 were compressed and the biomass increased with exposures under all
wavebands. Both the wavebands of UV-A + blue light (320—500 nm) and red light (600—700 nm) could ini-
tiate the spiral compression, growth and photosynthetic activities in A. platensis D-0083 efficiently. The effi-
ciencies per unit energy of irradiance to induce helix pitch changes in wavebands 320—500, 395—700,
510—700 and 610—700 nm were 0.070, 0.015, 0.021 and 0.045 μm/(W·m2) respectively. Waveband
320—500 nm had little suppression on effective quantum yield (Fv′/Fm′), electron transfer rate (ETR) and
phycocyanin (PC) fluorescence emission of the filaments but led to spiral compression and growth efficiently.
The waveband-dependent responses in spiral compression and specific growth rate appeared to be consistent
with the photosynthetic capability under the different light regimes.
Key words: Arthrospira platensis; Growth; Morphology; Photosynthetically active radiation (PAR); Photosynthesis
CLC number: Q948.11 Document code: A Article ID: 1000-3207(2016)03-0538-09
Light quality and intensity are critical environ-
mental signals that allow cells to sense the time of
day, the potential intracellular energy status, and to
tune metabolic activities towards its optimal growth
potential [1]. Rates of photosynthesis, growth, morpho-
genesis [2—7] and pigment composition [8—11] are some
of the light-dependent growth characteristics of cy-
anobacteria. Recently, blue light was suggested to
promote the metabolism of nitrogen-derived com-
pounds such as mycosporine-like amino acids
(MAAs), phycoerythrin and proteins in Porphyra leu-
costicta [12]. Blue light increases the chlorophyll and
phycocyanin contents and biomass production in
Spirulina fussiformis [13]. Light intensity is a major
trigger in the heterocyst and akinete differentiation of
many cyanobacterial species [7, 14—17].
The genes responsible for these light-sensitive
growth responses are well characterized and the sig-
naling mechanisms are reasonably well established [18].
In many cyanobacterial species, chromatic acclima-
tion (CA) results in lesser phycoerythrin (PE) and
more phycocyanin (PC) when grown under red than
in green light [19, 20]. The morphology of the Fremyella
diplosiphon is also affected during the CA process
when grown in red or green light [19, 21].
Cyanobacteria contain phytochrome, blue-light,
第 4 0 卷 第 3 期 水 生 生 物 学 报 Vol. 4 0, N o. 3
2 0 1 6 年 5 月 ACTA HYDROBIOLOGICA SINICA M a y, 2 0 1 6
Received date: 2015-05-11; Accepted date: 2015-12-04
Foundation item: Supported by the National Natural Science Foundation of China (No. 41430967, 41120164007, 31370381, 31170338);
Zhejiang Provincial Natural Science Foundation (No. LZ12C03001; LY14C030006); Project of Science and Technology
department of Zhejiang Province (No. 2015C33246)
Brief introduction of the author: Ma Zeng-Ling, Ph.D., Associate professor; E-mail: mazengling@wzu.edu.cn
Corresponding author: Gao Kun-Shan, E-mail: ksgao@xmu.edu.cn, Tel: 86-592-2187963
and UV-A receptors which can receive irradiance
spanning visible spectrum and near-UV [22-25]. Arth-
rospira platensis is an economically important cya-
nobacterium and commercially cultured around the
world to supply biomass and provide a rich source of
protein for the health food industry [26, 27]. Environ-
mental factors such as light and temperature deter-
mine the gross biomass productivity in Arthrospira
cultures [27]. Elevated photosynthetically active radi-
ation (PAR) levels decrease the helix pitch of A.
platensis [6], while UV radiation adding to PAR leads
to compressed spirals [28]. Oxidative stress induced by
light or UV stresses lead to broken trichomes in A.
platensis and A. variabilis PCC 7937[29, 30].
Although PAR is known to control the morpho-
logy and photosynthesis [6], we have yet to ascertain
the influence of the PAR region (i.e. 400 to 700 nm)
in the growth patterns, morphological and photosyn-
thetic changes in A. platensis. PAR penetration depth
decreased with wavelength in water body due to wa-
ter absorption generally increases with wavelen-
gth [31]. The filaments of Arthorspira spp. in natural
water body have to face the fluctuations of light qua-
lity and intensity when they stayed at different depth
in natural water body. Algal acclimation to fluctuat-
ing irradiance can lead to differently photosynthetic
rate, growth rates and cellular pigments content com-
pared to the cells acclimated to constant irradiance [32].
Therefore we predict that morphology, growth and
photosynthesis of A. platensis are waveband-specific
responding to solar PAR. Under this scenario, we in-
vestigated the morphology, growth and photosynthes-
is of A. platensis D-0083 filaments grown in quartz
tubes that were selectively exposed to various regions
of the PAR using three cut-off and one bandpass light
filters.
1 Materials and Methods
1.1 Experimental organism
Arthrospira platensis (D-0083) was obtained
from the Hainan Dainippon Ink and Chemicals (DIC)
microalgae CO. LTD. Hainan, China. A single
healthy spiral was chosen and all the trichomes were
propagated from it in a Zarrouk medium [33] containing
(g/L): NaHCO3—16.8, NaNO3—2.5, K2HPO4—0.5,
K2SO4—1.0, NaCl—1.0, MgSO4 .7H2O—0.2,
CaCl2—0.04, FeSO4 7H2O—0.01, EDTA—0.08 and
micronutrients. The cultures were aerated with
filtered (0.22 µm) air at 30℃ and 60 µmol/m2·s of
cool-white light (12 L∶12 D). Cells in the exponen-
tial growth phase were sampled at the beginning of
dark period and used in subsequent experiments.
1.2 Radiation treatments and measurement
The A. platensis cells were filtered, washed and
removed from the GF/C filter (25 mm Ø, Whatman)
and diluted with fresh Zarrouk medium to 0.16 opti-
cal densities at 560 nm (A560 nm). The cells were then
transferred to quartz tubes (2 cm Ø, length 12 cm) and
horizontally placed in opaque plastic containers with
removable and replaceable lids. The lids were fitted
with light filters for different wavebands (i.e. Ul-
traphan395 (395—700 nm), JB510 (510—700 nm),
HB610 (610—700 nm) and QB24 (320—500 nm)
that can be inserted and pulled out of the containers
upon demand to supply the A. platensis cells with the
appropriate radiation wavelengths (Fig. 1). The
loosened trichomes grown in laboratory, aerated with
filtered (0.22 µm) air at 30℃ and 60 µmol/(m2·s) of
cool-white light (12 L∶12 D), were set as the control
group, since the same irradiance level of full solar
spectrum would result in tightened spirals due to the
UV-stimulating effects [6].
The incident solar irradiance falling on the
quartz tube was measured by using a broadband EL-
Fig. 1 Irradiance spectrum of local solar radiation and the transmission characteristics of the Ultraphan 395 (395—700 nm), JB510
(510—700 nm), HB610 (610—700 nm) cut-off films and band-passing filter QB24 (320—500 nm) (A), and the corresponding doses (MJ) of
PAR, UV-A and UV-B during the period of 6 to 12 May, 2007 (B)
3 期 马增岭等: 钝顶螺旋藻形态、生长及光合作用对不同波段太阳辐射的响应 539
DONET filter radiometer (Real Time Computer,
Möhrendorf, Germany) equipped with three channels
dedicated for PAR (400—700 nm), UV-A (315—
400 nm) and UV-B (280—315 nm) wavebands. The
quartz containers were manually shaken for 4—5
times everyday to mix the culture and alleviate the ac-
cumulations of cells at the bottom of the tube [5]. Fur-
thermore and to minimize damage to cells caused by
the O2 or reactive oxygen species (ROS) accumula-
tion in the tubes triggered by photosynthetic activities
during the daytime [29], the cultures were aerated with
ambient air for 1min every night. Five replicates were
done for each treatment.
1.3 Temperature control
During the growth experiments, the containers
holding the quartz tubes were placed in a water bath
to maintain a (30±1)℃ temperature using a circula-
ting refrigerator (Eyela, CAP-3000, Tokyorikakikai
Co. Ltd. Tokyo, Japan).
1.4 Morphological examination
Morphological changes in A. platenssis spirals
were examined with a microscope (Zeiss Axioplan 2,
Carl Zeiss, Germany) after one week of growth
(13—19 April 2007) under different filters. Digital
images were recorded with a Zeiss Axicam and were
analyzed with an image analysis system (Axio Vision
3.0). Because the spirals of A. platensis D-0083 are
highly compressed, the helix pitch (the distance
between two neighboring spirals) was calculated as
the number of spirals per a given length. We ran-
domly estimated the helix pitch from at least 50 indi-
vidual filaments.
1.5 Determination of biomass and specific growth
rate
After the morphological examination of A.
platensis, cells in each tube were filtered to pre-dried
Whatman GF/C glass fiber filter (25 mm Ø, What-
man) and washed with 20 mL acidified distilled wa-
ter (pH 4) to remove residual salts. The cells were
then dried in an oven at 80℃ for 24h for the determ-
ination of biomass. Specific growth rate (µ, /d) of A.
platensis was calculated as µ = (lnx2-lnx1)/(t2-t1),
where x1 and x2 were the biomass at time t1 and t2, re-
spectively.
1.6 Determination of photosynthetic activity
To examine the effects of solar irradiance in dif-
ferent waveband on the photosynthetic capacity, the
effective photochemical quantum yield (Fv′/Fm′) and
elative electron transport (ETR) of cells grown under
different filters for a week were determined with a
portable pulse amplitude modulated fluoro- meter
(Water-PAM, Walz, Effeltrich, Germany). The acti-
nic light set at 100 μmol photons/(m2·s) and saturating
pulse was 5000 μmol photons/(m2·s) (0.8s). The ETR
was calculated as follows [34]: ETR [μmol e/(m2·s)] =
Fv′/Fm′ × 0.5 × PFD × A, where Fv′/Fm′ represents the
effective PSII quantum yield, PFD is the photosyn-
thetically active photon flux density, and A is the
fraction of incident photons absorbed by A. platensis
filaments[35]. The rapid light curves for ETR were
measured under eight different PAR levels (every
measurement lasted for 10s). The parameters of the
ETR curves were analyzed according to Webb, et
al. [36]: ETR = ETRmax × [1–e
(2α × E/ETRmax)], where α is
the efficiency of electron transport and E is the irradi-
ance. Five replicates were measured for each treat-
ment.
1.7 Measurement of chlorophyll fluorescence emis-
sion spectra
Arthorspira species contains chlorophyll a
(Chl.a), phycocyanobilin and allophycocyanin as light
harvesting pigments [37, 38]. In order to investigate
whether phycobilisome (PBS) was changed, we ex-
amined the changes in room-temperature chlorophyll
fluorescence of the cells grown under different filters
for a week. The chlorophyll fluorescence emission
spectra were measured with the spectrofluorimeter
(RF-5310PC, Shimadzu). The excitation wavelength
was set at 580 nm for PBS [29, 39].
1.8 Statistical analysis
A one-way ANOVA was used to analyze the dif-
ferences among treatments. When significant diffe-
rences occurred, Tukey’s HSD test was used to identi-
fy differences among treatment means. A confidence
level of 95% was used in all statistical analyses.
2 Results
The spectrum of local solar radiation and the
transmission of light intensities through the different
filters used in the radiation experiments were shown
in Fig. 1A. Transmitted light to solar radiation ratios
through Ultraphan 395 (395—700 nm), JB510
(510—700 nm), HB610 (610—700 nm) and QB24
(320—500 nm) were 284: 187: 88: 47 (Fig. 1A). The
sky was clear and provided with similar doses of so-
lar radiation throughout the duration of the experi-
ments (6—12 May 2007) (Fig. 1B).
Compared to the loosened filaments grown in
lab, the spirals of A. platensis D-0083 compressed
after one week exposure to solar irradiance in diffe-
rent wavebands (Fig. 2A). The length of trichomes in
various treatments was affected by the wavelength of
irradiance (Tab. 1). The length of filaments before ex-
posure (Control) and those cultured under QB24
ranged from 0 to 400 μm, but most (70%) of the fila-
ments under QB24 were shorter than 200 μm. No fila-
540 水 生 生 物 学 报 40 卷
ments longer than 300 μm were observed when cells
of A. platensis D-0083 were cultured outdoors under
Ultraphan395, JB510 and HB610 cut-off films. Fur-
thermore, the distribution of filaments with length
shorter than 100 mm followed the order Ultraphan395
(62%) > JB510 (46%) > HB610 (43%) > QB24
(21%) > Control (11%) indicating that more intense
irradiance doses led to shorter filaments. The helix
pitch of the spirals decreased significantly (P<0.01) in
cells grown under various cut-off films and band-pass
filter compared with the Control. There were signifi-
cant differences (P<0.05) in helix pitch between
means except in the comparison of JB510 with
HB610 treatments (Tab. 1, Fig. 2A).
When the variation in helix pitch of A. platensis
D-0083 was normalized to the energy that transmit-
ted through the filters, the efficiency of waveband
320—500 nm (i.e. UV-A + blue light or QB24) to
compress the spirals was much higher (P<0.01) than
other wavebands (Fig. 2B). The lowest effective irra-
diance to compress a spiral was the waveband cover-
ing fu l l v is ib le i r radiance (395—700 nm,
Ultraphan395). The efficiencies per unit energy of ir-
radiance to induce helix pitch changes in wavebands
320—500, 395—700, 510—700 and 610—700 nm
were 0.070, 0.015, 0.021 and 0.045 μm/(W·m2), re-
spectively (Fig. 2B).
Specific growth rate (μ, /d) of the cells after one
week exposure to different wavebands was highest
under Ultraphan395 and JB510, followed by HB610
and QB24 (Fig. 3A). It was 0.198 (± 0.021), 0.048 (±
0.007), 0.098 (± 0.007), 0.099 (± 0.003) and 0.090 (±
0.005) /d when grown indoors (Control) and covered
with QB24, Ultraphan395, JB510 and HB610 filters
during the exposure period (Fig. 3A).The efficiency
of luminous energy to influence the growth rate of A.
platensis D-0083 was similar to the variation in helix
pitch (Fig. 3B). Waveband-specific light efficiency to
induce specific growth rate variations for 320—500,
395—700, 510—700 and 610—700 nm were 0.0010
(± 0.0002), 0.0003 (± 0.0000), 0.0005 (± 0.0000) and
0.0010 (±0.000) /d/(W·m2), respectively.
Effective quantum yields (Fv′/Fm′) and electron
transfer rate [ETR, μmol e/(m2·s)] of A. platensis D-
0083 on the last noontime of exposure period (12
May, 2007) was highest under QB24 and lowest in
Ultraphan395 (Figs. 4A, B). Furthermore, quantum
yields and ETR increased with PAR wavebands to-
Fig. 2 Morphological changes (A) helix pitch change per energy [μm/(W·m2)] (B) of Arthrospira platensis D-0083 filaments under solar
exposures of different wavebands during the exposures (Fig. 1)
Scale bars in Fig. 2A indicate 100 μm (200 magnification. Means superscripted with different letters are significantly different (P<0.05).
Wavebands associated with each treatment are shown in Fig. 1A. The means and standard errors were based on 40 trichomes
Tab. 1 Distribution of trichome lengths and helix pitch of A. platensis D-0083 after one week of exposure under solar irradiance of the dif-
ferent wavebands
Treatments
Distribution of trichome lengths (%)
Helix pitch (μm)
0—100 mm 100—200 mm 200—300 mm 300—400 mm
Control 11 33 38 18 15.13±1.52a
QB24 21 50 25 4 11.81±0.77b
UL395 62 30 8 0 10.78±0.52c
JB510 46 41 13 0 11.29±0.51d
HB610 43 47 10 0 11.15±0.57d
Note: More than 100 trichomes in each treatment were randomly measured for the length determination. Means and standard er-
rors of helix pitch were calculated from at least 40 randomly measured trichomes. Means superscripted with different letters are sig-
nificantly different (P<0.05)
3 期 马增岭等: 钝顶螺旋藻形态、生长及光合作用对不同波段太阳辐射的响应 541
wards longer wavelength (UL395
and those under filter QB24, Ultraphan395, JB510
and HB610 were 0.50 (± 0.03), 0.51 (± 0.03), 0.18
(±0.03), 0.24 (± 0.04) and 0.31 (±0.04), respectively
(Fig. 4A). And the corresponding maximal electron
transfer rate (ETRmax) were 208.73 (± 2.13), 255.72 (±
7.98), 103.45 (± 1.66), 135.53 (± 7.09), 153.59 (±
7.50) μmol e/(m2·s), respectively (Fig. 4B). Com-
pared with the Control, the photosynthetic capability
(Fv′/Fm′ and ETR) was not (P>0.05) inhibited by the
irradiance between 320-500 nm but was significantly
(P<0.01) depressed by other wavebands.
When the cells grown under different filters were
excited at 580 nm, the emitted phycpcyanin (PC)
fluorescence intensity of the cells cultured indoors
(Control) and under QB24, Ultraphan395, JB510 and
HB610 filters were 144.5, 122.4, 149.8, 171.9 and
180.1, respectively (Fig. 5). Furthermore, compared
with the PC emission fluorescence of indoor cultures
(with peak at 646 nm), the emission peaks of cells
cultured under QB24, Ultraphan395, JB510 and
HB610 filters for a week shifted to longer wavelength
by 1, 7, 6 and 3 nm, respectively.
3 Discussion
Spirals of A. platensis D-0083 were compressed
and the biomass increased with exposures under diffe-
rent light wavebands. Both the wavebands of UV-A +
blue light (320—500 nm) and red light (600—700 nm)
could initiate the spiral compression, growth and pho-
tosynthetic activities in A. platensis D-0083 effi-
ciently. Our observations are in agreement with the
literatures with respect to the influence of various
wavebands to the growth and morphological regula-
tion of cyanobacteria. For example, cells of F. diplo-
siphon are long, brick-shaped and red under green
light, and smaller, spherical and blue-green under red
light due to synthesis of phycoerythrin or phycocyan-
in, respectively [19]. Furthermore, filaments of F. dip-
losiphon are shorter when grown in red light com-
pared to green light [19, 21]. Pure UV radiation seems
not capable of spiral modification in A. platensis [6].
However, the waveband spanning UV-A (320—400 nm)
to blue light (400—500 nm) could tighten the spirals
Fig. 3 Specific growth rate (/d) (A) and Specific growth rate per energy [/d/(W·m2)] (B) of Arthrospira platensis D-0083 exposed to differ-
ent wavebands of solar radiation from 6 to 12 May, 2007
Means superscripted with different letters are significantly different (P<0.05). Wavebands associated with each treatment are shown in Fig.
1A. The means and standard errors were based on five replicates
Fig. 4 Effective quantum yield (A) and relative electron transfer rate (B) at noontime of Arthrospira platensis D-0083 exposed to different
wavebands of solar radiation from 6 to 12 May, 2007
The means and standard errors were based on five replicates. Means superscripted with different letters are significantly different (P<0.05)
from each other. Please see Fig. 1A for the wavebands associated with each treatment
542 水 生 生 物 学 报 40 卷
of A. platensis D-0083 efficiently (Figs. 2A, B). It in-
dicates that blue light was more effective in trigger-
ing spiral compression in A. platensis, although any
irradiance with wavelength between 400 and 700 nm
might also induce the change. The similarity in spiral
helix pitch in JB510 and HB610 treatments and the
2X irradiance level under JB510 suggests that the
waveband 610—700 nm was a more effective trigger
to the spiral compression than 510—610 nm.
The specific growth rates of A. platensis D-0083
were similar when exposed to Ultraphan395 and
JB510, although the irradiance dose that the cells ac-
tually received was 1.5X in Ultraphan 395 compared
to JB510. This observation may mean that the maxi-
mal growth rate of the cells was not reached under fil-
ter JB510, HB610 and QB24 but was inhibited under
Ultraphan395. The lower specific growth rates ob-
served in this study than those under Ultraphan395
and aerated with ambient air [40] could be ascribed to
the build up of ROS produced from the photosynthe-
tic process [29, 30]. The cell depositions in the bottom of
quartz tubes could be attributed to the carbohydrate
accumulation during the photosynthesis [5] and the cell
deposits may have blocked the irradiance to reach the
entire photosynthetic cells of A. platensis D-0083.
Furthermore, the similar patterns in helix pitch and
specific growth rate (Figs. 2, 3) of A. platensis D-
0083 suggests the same irradiance waveband may be
responsible for morphological changes and growth
rate in Arthrospira species.
Cyanobacterial phytochrome (Cph) [41, 42] is si-
milar to plant phytochromes (red/farred photoreceptors)
that influence plant development including accessory
roles to sense the presence of UV-B and blue li-
ghts[43, 44]. Phycobilisome (PBS), the pigment-protein
complexes responsible for light harvesting in cy-
anobacteria, extend the absorption of light into red
and green regions of the visible spectrum to increase
energy capture for photosynthesis [20, 45]. The PBS
molds to light quality through CA depending on the
genetic characteristics of an organism, which in turn
is associated with its evolutionary environment [46, 47].
The increased intensities and red shifted (to longer
wavelengths) peaks of PC emission fluorescence (Fig.
5) under Ultraphan395, JB510 and HB610 revealed
the damage and structural modification of PBS in-
duced by the wavebands transmitted the filters [29, 39].
Furthermore, the more changes in PC emission fluor-
escence peaks of cells under filters Ultraphan395 and
JB510 compared with those under QB24 and HB610
indicated more structural modification of the PBS oc-
curred in the former. Another phytochrome-like
photoreceptor and regulator of CA in cynobac- teria is
RcaE that regulates light-dependent changes in phy-
cobiliprotein content [45, 48] and the cellular and fila-
ment morphology of F. diplosiphon [21]. However, the
exact molecular mechanism behind this morphologi-
cal regulation is unknown. In this study, the light
quality and intensity could not be clearly separated
due to the continuity and inhomogeneity of solar
spectrum as well as the flaw in transmissions of the
filters (Fig. 1A). Nevertheless, this study substanti-
ated that light quality has significant effects on mor-
phology and physiological activities of the filament-
ous cyanobacterium in that its spiral compression,
growth and photosynthetic activities were waveband-
specifically regulated.
Furthermore, Spirulina fussiformis when ex-
posed to blue light increased the production of C-phy-
cocyanin by photo-physiological mechanisms [13]
where the C-phycocyanin has high in vitro antioxi-
dant activity [49]. The effectiveness of blue and red
light to trigger the growth and morphological change
maybe related to the dominant absorbance of PC and
chlorophyll a (Chl. a) in the blue and red light re-
gions [50] or the regulatory role of Cph, PBS and RcaE
to promote cell development. As prokaryotic orga-
nism, the control of Cph, PBS and RcaE on morpho-
logy of cells or trichomes of A. platensis still needs to
be understood. However, the natural physiological
flexibility will undoubtedly facilitate the survival and
adaptation of an organism under rapidly changing en-
vironmental conditions including changes in spectral
component of solar radiation in natural water body.
4 Conclusion
In conclusion, the growth characteristics of A.
Fig. 5 Changes in fluorescence emission of phycocyanin (PC) in
A. platensis D-0083 cells grown under the exposures (Fig. 1). The
excitation wavelength 580 nm and the means were based on trip-
licate incubations
3 期 马增岭等: 钝顶螺旋藻形态、生长及光合作用对不同波段太阳辐射的响应 543
platensis D-0083 when exposed to solar irradiance re-
vealed the wavelength-dependent influence in
physiological and morphological regulations. Both
the waveband of UV-A + blue light (320—500 nm)
and the waveband of red light (600—700 nm) could
initiate the growth, spiral compression and photo-
synthetic activities in A. platensis D-0083 efficiently.
We speculated that the efficiency of visible light to in-
duce changes in morphology and growth of Arthrospira
spp. was related to the capabilities of wavelengths to
regulate the photosynthetic activities.
参 考 文 献:
Sharrock R A. The phytochrome red/far-red photoreceptor
superfamily [J]. Genome Biology, 2008, 9(8): 230
[1]
Dring M J. Photocontrol of development in algae [J]. Annual
Review of Plant Physiology and Plant Molecular Biology,
1998, 39: 157—174
[2]
Aguilera J, Gordillo F J L, Karsten U, et al. Light quality ef-
fect on photosynthesis and efficiency of carbon assimilation
in the red alga Porphyra leucosticte [J]. Journal of Plant
Physiology, 2000, 157(1): 86—92
[3]
Tsekos I, Niell F X, Aguilera J, et al. Ultrastructure of the
vegetative gametophytic cells of Porphyra leucosticta
(Rhodophyta) grown in red, blue and green light [J]. Phyco-
logical Research, 2002, 50(4): 251—264
[4]
Ma Z, Gao K. Photosynthetically active and UV radiation act
in an antagonistic way in regulating buoyancy of Arthros-
pira (Spirulina) platensis (cyanobacterium) [J]. Environ-
mental and Experimental Botany, 2009, 66(2): 265—269
[5]
Ma Z, Gao K. Photoregulation of morphological structure
and its physiological relevance in the cyanobacterium Arth-
rospira (Spirulina) platensis [J]. Planta, 2009, 230(2):
329—337
[6]
Singh S P, Montgomery B. Determining cell shape: adaptive
regulation of cyanobacterial cellular differentiation and mor-
phology [J]. Trends in Microbiology, 2011, 19(6): 278—285
[7]
Babu T S, Kumar A, Varma A K. Effect of light quality on
phycobilisome components of the cyanobacterium Spirulina
platensis [J]. Plant Physiology, 1991, 195(2): 492—497
[8]
Takano H, Arai T, Hirano M, et al. Effects of intensity and
quality of light on phycocyanin production by a marine cy-
anobacterium Synechococcus sp. NKBG 042902 [J]. Ap-
plied Microbiology and Biotechnology, 1995, 43(6):
1014—1018
[9]
Tandeau de Marsac N. Phycobiliproteins and phycobili-
somes: the early observations [J]. Photosynthesis Research,
2003, 76(1): 197—205
[10]
Vijaya V, Anand N. Blue light enhance the pigment synthe-
sis in cyanobacterium Anabaena ambigua Rao (Nostacales)
[J]. Archive of ARPN Journal of Agricultural and Biological
Science, 2009, 4(3): 36—43
[11]
Korbee N, Figueroa F L, Aguilera J. Effect of light quality
on the accumulation of photosynthetic pigments, proteins
and mycosporine-like amino acids in the red alga Porphyra
leucosticta (Bangiales, Rhodophyta) [J]. Journal of Photo-
chemistry and Photobiology B: Biology, 2005, 80(2): 71—78
[12]
Madhyastha H K, Vatsala T M. Pigment production in
spirulina fussiformis in different photophysical conditions
[J]. Biomolecular Engineering, 2007, 24(3): 301—308
[13]
Adams D G, Duggan P A. Tansley review No. 107. hetero-
cyst and akinete differentiation in cyanobacteria [J]. New
Phytologist, 1999, 144(1): 3—33
[14]
Moore D, Odonohue M, Garnett C, et al. Factors affecting
akinete differentiation in Cylindrospermopsis raciborskii
(Nostocales, Cyanobacteria) [J]. Freshwater Biology, 2005,
50(2): 345—352
[15]
Thompson P A, Jamesson I, Blackburn S I. The influence of
light quality on akinete formation and germination in the to-
xic cyanobacterium Anabaena circinalis [J]. Harmful Algae,
2009, 8(3): 504—512
[16]
Kaplan-Levy R, Hadas O, Summers M L, et al. Akinetes:
dormant cells of cyanobacteria. In: Lubzens E, Cerda J,
Clark M (Eds.), Dormancy and Resistance in Harsh Environ-
ments [M]. Springer, Berlin/Heidelberg. 2010, 5—27
[17]
Grossman A R, Bhaya D, He Q. Tracking the light environ-
ment by cyanobacteria and the dynamic nature of light har-
vesting [J]. Journal of Biological Chemistry, 2001, 276(15):
11449—11452
[18]
Bennett A, Bogorad L. Complementary chromatic adapta-
tion in a filamentous blue-green alga [J]. Journal of Cell Bio-
logy, 1973, 58: 419—435
[19]
Kehoe D M, Gutu A. Responding to color: The regulation of
complementary chromatic adaptation [J]. Annual Review of
Plant Biology, 2006, 57(1): 127—150
[20]
Bordowitz J R, Montgomery B L. Photoregulation of cellu-
lar morphology during complementary chromatic adaptation
requires sensor-kinase-class protein RcaE in Fremyella dip-
losiphon [J]. Journal of Bacteriology, 2008, 190(11):
4069—4074
[21]
Tsinoremas N F, Schaefer M, Golden S S. Blue and red light
reversibly control psbA expression in the cyanobacterium
Synechococcus sp strain PCC 7942 [J]. Journal of Biologi-
cal Chemistry, 1994, 269(23): 16143—16147
[22]
Lamparter T, Mittmann F, Gärtner W, et al. Characteriza-
tion of recombinant phytochrome from the cyanobacterium
Synechocystis [J]. Proceedings of the National Academy of
Sciences of the United States of America, 1997, 94(22):
11792—11797
[23]
Alfonso M, Perewoska I, Kirilovsky D. Redox control of ps-
bA gene expression in the cyanobacterium Synechocystis
PCC 6803: involvement of the cytochrome b6/f complex [J].
Plant Physiology, 2000, 122(2): 505—515
[24]
Hirose Y, Rockwell N C, Martin S S, et al. Green/red cya-
nobacteriochromes regulate complementary chromatic accli-
[25]
544 水 生 生 物 学 报 40 卷
mation via a protochromic photocycle [J]. Proceedings of the
National Academy of Sciences of the United States of Ameri-
ca, 2013, 110(13): 4974—4979
Torzillo G, Vonshak A. Biotechnology of algal mass cultiva-
tion. In: Fingerman M, Nagabhushanam R (Eds.), Recent ad-
vances in marine biotechnology, Biomaterials and biopro-
cessing [M]. Science Publishers, Inc, Plymouth. 2003,
45—77
[26]
Sili C, Torzillo G, Vonshak A. Arthrospira (Spirulina). In:
Whitton B A (Eds.), Ecology of Cyanobacteria II: their Di-
versity in Space and Time [M]. Springer. 2012, 677—705
[27]
Wu H, Gao K, Villafañe V, et al. Effects of solar UV radia-
tion on morphology and photosynthesis of the filamentous
cyanobacterium Arthrospira platensis [J]. Applied Environ-
mental Microbiology, 2005, 71(9): 5004—5013
[28]
Ma Z, Gao K. Spiral breakage and photoinhibition of Arth-
rospira platensis (Cyanophyta) caused by accumulation of
reactive oxygen species under solar radiation [J]. Environ-
mental and Experimental Botany, 2010, 68(2): 208—213
[29]
Rastogi R P, Singh S P, Hader D P, et al. Detection of reac-
tive oxygen species (ROS) by the oxidant-sensing probe 2′,
7′-dichlorodihydrofluorescein diacetate in the cyanobacte-
rium Anabaena variabilis PCC 7937 [J]. Biochemical and
Biophysical Research Communications, 2010, 397(3):
603—607
[30]
Kokhanoovsky A A. The depth of sunlight penetration in
cloud fields for remote sensing [J]. IEEE Geoscience and Re-
mote Sensing Letters, 2004, 1(4): 242—245
[31]
van de Poll W H, Visser R J W, Buma A G. Acclimation to a
dynamic irradiance regime changes excessive irradiance
sensitivity of Emiliania huxleyi and Thalassiosira weissflo-
gii [J]. Limnology and Oceanography, 2007, 52(4):
1430—1438
[32]
Zarrouk C. Contribution a letude d une cyanophycee. Influ-
ence de diverse facteurs physiques et chimiques sur la crois-
sance et la photosynthese de Spirulina maxima (Setch et
Gardner) Geitler. Ph. D. Thesis, University of Paris, France.
1966
[33]
Genty B, Briantais J M, Baker N R. The relationship
between the quantum yield of photosynthetic electron trans-
port and quenching of chlorophyll fluorescence [J]. BBA
General Subjects, 1989, 990(1): 87—92
[34]
Franklin L A, Badger M R. A comparison of photosynthetic
electron transport rates in macroalgae measured by pulse
amplitude modulated chlorophyll fluorometry and mass
spectrometry [J]. Journal of Phycology, 2001, 37(5):
756—767
[35]
Webb W L, Newton M, Starr D. Carbon dioxide exchange of
Alnus rubra: mathematical model [J]. Oecologia, 1974,
17(4): 281—291
[36]
Cohen Z. The chemicals of Spirulina. In: Vonshak A (Eds.),[37]
Spirulina platensis (Arthrospira): Physiology, Cell-biology
and Biotechnology [M]. Taylor ↦ Francis Publishers,
London. 1997, 175—204
Nuhu A A. Spirulina (Arthrospira): an important source of
nutritional and medicinal compounds [J]. Journal of Marine
Biology, 2013: Article ID 325636, 8
[38]
Wen X, Gong H, Lu C. Heat stress induces an inhibition of
excitation energy transfer from phycobilisomes to photosys-
tem II but not to photosystem I in a cyanobacterium
Spirulina platensis [J]. Plant Physiology and Biochemistry,
2005, 43(4): 389—395
[39]
Gao K, Ma Z. Photosynthesis and growth of Arthrospira
(Spirulina) platensis (Cyanophyta) in response to solar UV
radiation, with special reference to its minor variant [J]. En-
vironmental and Experimental Botany, 2008, 63(1—3):
123—129
[40]
Hughes J, Lamparter T, Mittman F, et al. A prokaryotic
phytochrome [J]. Nature, 1997, 386(6626): 663
[41]
Yeh K C, Wu S H, Murphy J T, et al. A cyanobacterial
phytochrome two-component light sensory system [J]. Sci-
ence, 1997, 277(5331): 1505—1508
[42]
Whitelam G C, Devlin P F. Roles of different phytochromes
in Arabidopsis photomorphogenesis [J]. Plant, Cell Environ-
ment, 1997, 20(6): 752—758
[43]
Kim B C, Tennessen D J, Last R L. UV-B-induced photo-
morphogenesis in Arabidopsis thaliana [J]. Plant Journal,
1998, 15(5): 667—674
[44]
Kehoe D M, Grossman A R, The molecular mechanisms
controlling complementary chromatic adaptation. In: Pes-
chek G A, Löffelhardt W, Schmetterer G (Eds.), The Photo-
trophic Prokaryotes [M]. Kluwer Academic/Plenum Publi-
shers, New York. 1999, 61—69
[45]
Montgomery B L. Sensing the light: photoreceptive systems
and signal transduction in cyanobacteria [J]. Molecular Mi-
crobiology, 2007, 64(1): 16—27
[46]
Gutu A, Kehoe D M. Emerging perspectives on the mecha-
nisms, regulation, and distribution of light color acclimation
in cyanobacteria [J]. Molecular Plant, 2012, 5(1): 1—13
[47]
Terauchi K, Montgomery B L, Grossman A R, et al. RcaE is
a complementary chromatic adaptation photoreceptor re-
quired for green and red light responsiveness [J]. Molecular
Microbiology, 2004, 51(2): 567—577
[48]
Madhyastha H K, Sivashankari S, Vatsala T M. C-phycoc-
yanin from Spirulina fussiformis exposed to blue light
demonstrates higher efficacy of in vitro antioxidant activity
[J]. Biochemical Engineering Journal, 2009, 43(2):
221—224
[49]
Minkova K M, Tchernov A A, Tchorbadjieva M I, et al.
Purification of C-phycocyanin from Spirulina (Arthrospira)
Fusiformis [J]. Journal of Biotechnology, 2003, 102(1):
55—59
[50]
3 期 马增岭等: 钝顶螺旋藻形态、生长及光合作用对不同波段太阳辐射的响应 545
钝顶螺旋藻形态、生长及光合作用对不同波段太阳辐射的响应
马增岭1 M. Arocena Joselito1, 2 高坤山3
(1. 浙江省亚热带水环境与海洋生物资源保护重点实验室, 温州大学, 温州 325035; 2. 北英属哥伦比亚大学环境科学与工程学
院, 乔治王子城 V2N4Z9; 3. 近海海洋环境科学国家重点实验室, 厦门大学, 厦门 361005)
摘要: 为探讨中不同波段的光合有效辐射对钝顶螺旋藻(Arthrospira platensis)形态、生长及光合作用的影响,
实验将钝顶螺旋藻D-0083藻液转入带塞的石英管中, 石英管水平置于阳光下并在其上覆盖不同的截止型和带
通型滤光片, 以使藻丝接受不同波段的太阳辐射; 并检测其生长、形态与光合活动的变化。结果发现: 所有波
段 (320—500、395—700、510—700和610—700 nm) 光合有效辐射下的藻丝均螺旋变紧且生物量增加。其
中以包含少量紫外辐射A (Ultraviolet-A)的蓝光波段 (320—500 nm)和红光波段(600—700 nm) 对藻丝形态变
化、生长及光合速率的诱发效率较高。在320—500、395—700、510—700和 610—700 nm波段上的单位能
量光照引起钝顶螺旋藻螺距变化的效率分别为0.070、0.015、0.021、0.045 μm/(W·m2)。波段320—500 nm
虽然会轻微抑制钝顶螺旋藻D-0083的有效光化学效率(Fv′/Fm′)、电子传递速率(ETR)和藻蓝蛋白的荧光发射,
但是却能够有效诱导其藻丝变紧促进生长。此外, 钝顶螺旋藻D-0083的藻丝变紧程度、比生长速率变化与不
同波段太阳辐射下藻丝体的光合性能相一致。该研究表明任何波段的光合有效辐射都能使螺旋藻藻丝螺旋
变紧并引发生长和光合作用, 其中以蓝光和红光的效率最高。
关键词: 钝顶节旋藻; 生长; 形态; 光合有效辐射(PAR); 光合作用
546 水 生 生 物 学 报 40 卷