环境因子节律性变动对潮间带大型海藻孢子萌发、早期发育和生长的影响及其生理生态学机制
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摘要
一、温度日节律波动对潮间带大型海藻孢子萌发、早期发育、生长、生化组成、营养吸收和光合作用的影响
1.温度日节律波动对孔石莼生长和生化组成的影响
实验共设5个温度日节律波动处理(20±2℃,20±4℃,20±6℃,20±8℃,20±10℃)和1个恒温(20℃±0℃)对照。实验结果表明:每天不同温度波动幅度对孔石莼(Ulva pertusa)的生长率具有明显的影响(P<0.05),其中(20±2)℃、(20±4)℃和(20±6)℃与恒温对照相比能明显促进孔石莼的生长,(20±10)℃则显著抑制孔石莼的生长;孔石莼相对生长率(RGR)与温度波动幅度(TA)的关系符合如下多项式方程: G=β_0+β_1(TA)+β_2(TA)~2其中G代表相对生长率,TA代表温度波动幅度,β0为方程在G轴上的截距,β_1和β2为回归系数。当平均温度为20℃时,对孔石莼生长最有利的温度波动幅度为±3.69℃。温度日节律波动对孔石莼Chl-a、Chl-b、蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05),(20±2)℃和(20±4)℃处理组的Chl-a含量显著高于恒温对照组,(20±2)℃、(20±4)和(20±6)℃处理的蛋白质和可溶性糖的绝对含量明显高于恒温对照组,但当温度波动幅度大于±8℃时,可溶性糖和细胞防御物质游离脯氨酸的含量明显增加(P<0.05)。
2.温度日节律波动对带形蜈蚣藻四分孢子放散、孢子萌发和早期发育的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。观察了带形蜈蚣藻孢子的放散,并在不同温度条件下测定了带形蜈蚣藻孢子萌发率和早期发育。实验结果表明带形蜈蚣藻四分孢子囊携四个孢子同时从藻体中排出,孢子囊膜破裂后四个孢子同时放散;四分孢子的萌发类型为Nemalion type;从带形蜈蚣藻四分孢子附着至第三次分裂,藻体的大小没有发生明显的变化;适宜的温度波动促进了带形蜈蚣藻四分孢子的萌发和早期发育。
3.温度日节律波动对真江蓠生长和营养吸收的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。研究了真江蓠在不同温度条件下的生长、元素积累和营养吸收。实验结果表明温度日节律波动能够促进了真江蓠的生长、对碳元素和氮元素的积累以及对水体中NO3--N的吸收和移除。
4.温度日节律波动中昼高温、夜低温对真江蓠营养吸收和光合作用的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。研究了在不同温度条件下昼高温和夜低温对真江蓠的营养吸收和光合作用。实验结果表明温度日节律波动中昼高温不但促进了真江蓠对水体中NO_3~--N的吸收,也促进了其净光合作用速率;同时在夜低温能够抑制真江蓠的夜间呼吸作用。上述结论能够解释温度日节律波动促进潮间带海藻生长的现象。
二、节律性干出对孔石莼生长和生化组成的影响
实验共设7个节律性干出的处理,即每12h干出0.0h、0.5h、1.0h、2.0h、3.0h、4.0h和5.0h。实验结果表明:每12h不同干出时间对孔石莼(Ulva pertusa)的生长具有明显的影响(P<0.05),其中干出0.5h、1.0h和2.0h与不干出处理相比能明显促进孔石莼的生长,干出5h则显著抑制孔石莼的生长(P<0.05);节律性干出对孔石莼叶绿素a、叶绿素b、蛋白质、可溶性糖和游离脯氨酸含量具有明显的影响(P<0.05),干出0.5h、1.0h和2.0h处理的可溶性糖的绝对含量明显高于不干出处理,干出时间过长(3.0h、4.0h和5.0h)不但使孔石莼生长物质的含量减少,并且使孔石莼的渗透溶质增加,当干出时间为5h时,可溶性糖含量突然增高;游离脯氨酸的含量也随着干出时间的增加而增加。
三、盐度日节律波动对孔石莼生长和生化组成的影响
实验共设5个盐度日节律变化处理(30±3、30±6、30±9、30±12和30±15)和一个恒盐(30±0)对照处理。实验结果表明:孔石莼能够耐受较大范围的盐度日节律性连续变化,这种变化虽然对孔石莼的生长不利,但是仍然能够使孔石莼的相对生长率保持在12%以上;每天不同盐度变化对孔石莼的生长率具有明显的影响(P<0.05),盐度变化幅度±3~±15与恒盐对照相比明显不利于孔石莼的生长,盐度变化幅度越大,孔石莼生长越缓慢。盐度日节律连续变化对孔石莼蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05)。同恒盐处理相比,盐度日节律连续变化对孔石莼叶绿素的含量的影响不明显,但叶绿素a和叶绿素b含量都有随着盐度变化幅度的增大而逐渐减少的趋势。盐度日节律性连续变化使孔石莼蛋白质和可溶性糖的相对含量显著增加(P<0.05),随着盐度变化幅度的增大,蛋白质和可溶性糖增加的趋势越明显,并且在盐度日节律性连续变化下,蛋白质增加的速度比可溶性糖增加的速度要快。同时,游离脯氨酸的相对含量随着盐度变化幅度的增加而增加,但是在缓和的盐度日节律性连续变化下(30±3)孔石莼游离脯氨酸的含量却显著小于恒盐对照组(P<0.05)。
四、光对孔石莼生长和生化组成的影响
1.光照强度对孔石莼生长和生化组成的影响
研究了50～400μmol/(m2·s)范围内光照强度对孔石莼生长和藻体生化组成的影响。研究结果表明光照强度对孔石莼的生长率具有明显的影响(P<0.05),孔石莼在光照强度为250μmol/(m~2·s)时有最大的生物量积累。光照强度低于250μmol/(m~2·s)时,孔石莼的生长率有随着光照强度的降低而减小的趋势;光照强度高于250μmol/(m~2·s)时,孔石莼的生长率随着光照强度的升高而降低(P<0.05);过高或过低的光照强度均不利于孔石莼的生长,光照强度在100μmol/(m~2·s)以下时,孔石莼的生长明显受到不利的影响,光照强度至350μmol/(m~2·s)以上、实验超过6天后藻体叶片产生溃斑,并产生了明显的光抑制。不同光照强度对孔石莼的叶绿素、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05)。随着光照强度的增加,孔石莼的叶绿素a和叶绿素b的含量逐渐变小;低光照条件可促进了孔石莼蛋白质的积累;当光照强度超过350μmol/(m~2·s)时,游离脯氨酸的含量明显增加。光照强度对孔石莼化学组成也具有明显的影响(P<0.05)。随着光照强度的升高,藻体的N元素含量有所升高,但光强达到350μmol/(m~2·s)后,随着光强的增加,N元素含量反而降低;C元素的含量在光照强度为50~350μmol/(m~2·s)范围内变化不明显,但当光照强度达到350μmol/(m~2·s)后,C元素的含量明显升高;H元素的含量在50~400μmol/(m~2·s)范围内,具有随着光照强度增加而增加的趋势。
2.光照强度日节律波动对孔石莼生长和生化组成的影响
本研究由两个实验组成。实验1共设5个光照强度波动幅度处理( 200±40μmol/(m~2·s)、200±80μmol/(m~2·s)、200±120μmol/(m~2·s)、200±160μmol/(m~2·s)和200±180μmol/(m~2·s))和1个恒定光照强度200μmol/(m~2·s)对照。实验1的结果表明:每天不同的光照强度波动幅度对孔石莼的生长率具有明显的影响( P<0.05 ),其中200±40μmol/(m~2·s)、200±80μmol/(m~2·s)和200±120μmol/(m~2·s)与恒定光照强度200μmol/(m~2·s)对照相比能明显促进孔石莼的生长;光照强度日节律波动对孔石莼Chl-a、Chl-b、蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05),光照波动幅度为80μmol/(m~2·s)处理下孔石莼的叶绿素a和叶绿素b的含量显著高于波动幅度为120μmol/(m~2·s)、160μmol/(m~2·s)和180μmol/(m~2·s)的处理(P<0.05),但恒定光照强度处理与光照强度波动处理下孔石莼叶绿素a和叶绿素b的含量相比没有显著的差异(P>0.05);孔石莼的蛋白质和可溶性糖含量具有随着光照强度波动幅度增大而减小的趋势。游离脯氨酸含量以恒定光照强度下最高,显著高于其余各光照强度波动处理(P<0.05)。实验2共设3个光照强度波动幅度处理(250±50μmol/(m~2·s)、250±100μmol/(m~2·s)和250±150μmol/(m~2·s))和一个恒定光照强度250μmol/(m~2·s)对照。实验2的结果表明:每天不同的光照强度波动幅度对孔石莼的生长率具有明显的影响( P<0.05 ), 250±50μmol/(m~2·s)、250±100μmol/(m~2·s)和250±150μmol/(m~2·s)处理下孔石莼终生物量和相对生长率都显著高于恒定光照强度200μmol/(m~2·s)对照组(P<0.05);恒定光照强度250μmol/(m~2·s)下孔石莼叶绿素a的含量显著小于波动幅度为50μmol/(m~2·s)和100μmol/(m~2·s)的处理(P<0.05),但与波动幅度为150μmol/(m~2·s)处理下孔石莼叶绿素a的含量相比差异不显著(P>0.05);恒定光照强度250μmol/(m~2·s)处理与光照强度波动处理下孔石莼叶绿素b的含量没有显著的差异(P>0.05)。光照波动幅度为50μmol/(m~2·s)处理下孔石莼叶绿素b的含量显著高于波动幅度为150μmol/(m~2·s)的处理(P<0.05);孔石莼的蛋白质和可溶性糖含量在恒定光照强度下都比在光照强度波动处理下高;游离脯氨酸含量以恒定光照强度下最高,显著高于其余各光照强度波动处理(P<0.05)。
3.不同光照强度下光暗周期对孔石莼生长和生化组成的影响
研究了不同光照强度下光暗周期对孔石莼(Ulva pertusa)生长和生化组成的影响。发现在长日照能够明显促进孔石莼的生长,在全光照和全黑暗处理对孔石莼的生长不利。实验1光照强度为110μmol/(m~2·s),实验2光照强度为240μmol/(m~2·s),两种光照强度下分别设置11个光暗周期处理(L:D=24:0、20:4、16:8、14:10、13:11、12:12、11:13、10:14、8:16、4:20和0:24)。实验结果表明:不同光暗周期对孔石莼的生长率具有明显的影响(P<0.05),除全光照处理外,孔石莼的生物量具有随着光照时间的增加而增加的趋势;随着日照时间的减少,叶绿素a和叶绿素b的含量都有逐渐下降的趋势;蛋白质和可溶性糖的含量在极长和极短日照长度处理中都有增加的趋势;随着光照时间的增加,游离脯氨酸的含量也逐渐增加。
4.光谱成分及其变动对孔石莼生长和生化组成的影响
研究了光谱成分及其波动对孔石莼(Ulva pertusa)生长和生化组成的影响。实验共设11个光谱成分及其波动的处理(白光、橙色光、绿色光、红色光、蓝色光、橙色光:绿色光=6:6-h、橙色光:红色光=6:6-h、绿色光:红色光=6:6-h、绿色光:蓝色光=6:6-h、红色光:蓝色光=6:6-h、蓝色光:橙色光=6:6-h)。实验结果表明:橙色光、蓝色光同白光相比对孔石莼的生长没有明显的影响,而红色光和绿色光对孔石莼的生长产生了明显的抑制,尤其以绿色光的抑制作用最为显著(P<0.05)。光谱变化处理中,绿色光:红色光=6:6-h处理出现了光增益效应。红色光和蓝色光下孔石莼的叶绿素a和叶绿素b的含量显著小于白光处理(P<0.05)。单色光的波动有利于增加孔石莼叶绿素、蛋白质和可溶性糖的含量,并能减弱游离脯氨酸在藻体内的积累。
ⅠEffects of diel fluctuating temperature on germination, growth, biochemical composition, nutrient uptake and photosynthesis of several macroalgae
1 Effects of circadian rhythms of fluctuating temperature on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under (20±2)℃, (20±4)℃, (20±6)℃, (20±8)℃and (20±10)℃circadian rhythms of fluctuating temperature conditions, and constant temperature of 20℃was used as the control. The growth rate of macroalga at (20±2)℃, (20±4)℃and (20±6)℃were significantly higher than that at constant temperature of 20℃, while growth rate at (20±8)℃and (20±10)℃were significantly lower than that at constant temperature of 20℃. The growth rate of macroalga was a quadratic function of the thermal amplitude. Such a growth model can be described by G=β_0+β_1(TA)+β_2(TA)_2 where G represents the relative growth rate, TA is thermal amplitude in degree Celsius,β0 is the intercept on the G axis, andβ_1 andβ_2 are the regression coefficients. The optimal thermal amplitude for the growth of thallus at mean temperature of 20℃was estimated to be±3.69℃. Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating temperature caused various influences (P<0.05).The content of chlorophyll, protein and total solute carbohydrate at (20±2)℃ and (20±4)℃were slightly higher than those at constant temperature of 20℃, however no statistically significant differences were found among them (P>0.05). While osmolytes (total solute carbohydrate and free proline) at (20±10)℃were significantly higher than that at 20℃(P<0.05). Therefore, more chlorophyll and carbohydrate production might account for the enhancement in the growth of macroalga at the diel fluctuating temperatures in the present study.
2 Effects of diel fluctuating temperature on germination and early growth of Grateloupia turuturu
The marcoalga Grateloupia turuturu was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The spores germination, early growth were observed in this experiment. The type of G. turutur germination belongs to Nemalion type. There was no significant difference in the spores’size between 2 cells stage and 8 cells stage after spores attached(P>0.05). The germination rate of macroalga at (15±3)℃, (20±3)℃were significantly higher than that at constant temperature of 15℃and 20℃respectively(P<0.05).
3 Effects of diel fluctuating temperature on growth and nutrient uptake of Gracilaria vermiculophylla
The marcoalga Gracilaria vermiculophylla was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The specific growth rate, nutrient uptake and NO3--N removal were determined in this experiment. The result showed that diel fluctuating temperature can enhance the growth rate, nutrient uptake and NO3--N removal of Gracilaria vermiculophylla than that under the constant temperature(P<0.05).
4 Effects of high day temperature and low night temperature on nutrient uptake and photosynthesis of Gracilaria vermiculophylla
The marcoalga Gracilaria vermiculophylla was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The specific growth rate, nutrient uptake and photosynthetic rate and respiratory rate were determined in this experiment. The results showed that high day temperature and low night temperature play the different role during the temperature fluctuation. High day temperature not only can accelerate the NO3--N removal of Gracilaria vermiculophylla but also can enhance the net photosynthetic rate(P<0.05), and low night temperature can inhibit the respiration compare with the macroalga under the constant temperature(P<0.05).
ⅡEffects of periodical emersion on growth and biochemical composition of Ulva pertusa
The influence of emersion on the growth and biochemical composition were investigated with Ulva pertusa thalli grown under laboratory conditions with periodical emersion and submersion treatments. The accumulated biomass and daily increments of thalli with 0.5h, 1.0h and 2.0h periodical emersion every 12h were faster than those without emersion, while growth rate with 5h periodical emersion every 12h was significantly lower than that without emersion. Analysis of biochemical composition at the final stages of thalli growth revealed that periodical emersion caused various influences (P<0.05). The final content of total solute carbohydrate with 0.5h, 1.0h and 2.0h periodical emersion every 12h was significantly higher than that without emersion, while osmolyte with 5h periodical emersion every 12h was significantly higher than that without emersion (P<0.05). Therefore, mild periodical emersion could enhance the growth of Ulva pertusa, while severe periodical emersion could inhibit the growth of Ulva pertusa.
ⅢEffects of circadian rhythms of fluctuating salinity on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva Pertusa was cultured under 30±3, 30±6, 30±9, 30±12 and 30±15 circadian rhythms of fluctuating salinity conditions, and constant salinity of 30 was used as the control. The growth rate of macroalga at fluctuating salinity conditions were significantly lower than that at constant salinity of 30, while the SGR of U. pertusa at different fluctuating salinity conditions were all kept at least 12%. Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating salinity caused various influences (P<0.05). The content of chlorophyll a and chlorophyll b at fluctuating salinity conditions were slightly lower than those at constant salinity of 30, however no statistically significant differences were found among them (P>0.05). The content of protein and total solute carbohydrate at fluctuating salinity conditions were significantly higher than those at constant salinity of 30 (P<0.05), and there was a trend that the content of proline increased with the increase of amplitude of fluctuating salinity, while the content of proline at mild change of salinity (30±3) was significantly lower than that at constant salinity (P<0.05).
ⅣEffects of light on growth and biochemical composition of Ulva pertusa
1. Effects of light intensity on growth and chemical constituents of Ulva pertusa The influence of light on the growth and chemical constituents were investigated with Ulva pertusa thalli growth under laboratory conditions with different light intensity. There was a trend that the growth rate of macroalga decreased with the decrease of light intensity under 250μmol/(m~2·s), but when light intensity exceed 250μmol/(m~2·s), the growth rate of macroalga also decrased with the increase of light intensity(P<0.05). Analysis of chemical constituents at the final stages of thalli growth revealed that different light intensity caused various influences (P<0.05). There was a trend that the content of chlorophyll a and chlorophyll b decreased with the increase of light intensity. The synthesization of protein was accelerated within the lower light intensity. While osmolytes (free proline) at 350μmol/(m~2·s) and 400μmol/(m~2·s) was significantly higher than other treatments (P<0.05). The content of nitrogen increased with the increase of light intensity from 50 to 300μmol/(m~2·s), while the content of nitrogen decreased with the increase of light intensity when light intensity exceed 350μmol/(m~2·s), the content of carbon has the same trend with the trend of nitrogen, and the content of hydrogen increased with the increase of light intensity in this study.
2. Effects of circadian rhythms of fluctuating light intensity on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under 200±40μmol/(m~2·s), 200±80μmol/(m~2·s), 200±120μmol/(m~2·s), 200±160μmol/(m~2·s) and 200±180μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions, and constant light intensity of 200μmol/(m~2·s) was used as the control in experiment 1. The growth rate of macroalga at 200±40μmol/(m~2·s), 200±80μmol/(m~2·s) and 200±120μmol/(m~2·s) were significantly higher than that at constant light intensity of 200μmol/(m~2·s) (P<0.05). Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating light intensity caused various influences (P<0.05).While osmolytes (free proline) at constant light intensity of 200μmol/(m~2·s) was significantly higher than those at other treatment (P<0.05).The marcoalga Ulva pertusa was cultured under 250±50μmol/(m~2·s), 250±100μmol/(m~2·s) and 250±150μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions, and constant light intensity of 250μmol/(m~2·s) was used as the control in experiment 2. The growth rate of macroalga at 250±50μmol/(m~2·s), 250±100μmol/(m~2·s) and 250±150μmol/(m~2·s) were significantly higher than that at constant light intensity of 250μmol/(m~2·s) (P<0.05). Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating light intensity caused various influences (P<0.05). .The content of chlorophyll-a at 250±50μmol/(m~2·s) and 250±100μmol/(m~2·s) was significantly higher than that at constant light intensity of 250μmol/(m~2·s) (P<0.05), the content of protein and total solute carbohydrate at 250±50μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions were significantly higher than those at constant temperature of 250μmol/(m~2·s), osmolytes (free proline) at constant light intensity of 250μmol/(m~2·s) was significantly higher than those at other treatment (P<0.05).
3. Effects of light-dark cycles on growth and biochemical composition of Ulva pertusa under different light intensity conditions
The marcoalga Ulva pertusa was cultured under L:D=24:0、20:4、16:8、14:10、13:11、12:12、11:13、10:14、8:16、4:20 and 0:24 light-dark cycles under 110μmol/(m~2·s) and 240μmol/(m~2·s) light intensity conditions, respectively. There was a trend that the growth rates of macroalga decreased with the increase of light period. The synthesization of protein and total carbohydrate were accelerated within the shorter light period and longer light period. And free proline was increased with the increase of light period.
4. Effects of light spectra and fluctuating of light spectra on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under White, Yellow, Green, Red, Blue, Yellow: Green=6:6h, Yellow: Red=6:6h, Green: Red=6:6h, Green: Blue=6:6h, Red: Blue=6:6h, Blue: Yellow=6:6h light spectra conditions. The growth rate of macroalga at Red light and Green light were significantly lower than that at White light. Emerson enhancement effect occurred at Green: Red=6:6h treatment. The content of chlorophyll, protein and total solute carbohydrate was accelerated by the fluctuating of light spectra. The content of praline was alleviated by the fluctuating of light spectra
1.温度日节律波动对孔石莼生长和生化组成的影响
实验共设5个温度日节律波动处理(20±2℃,20±4℃,20±6℃,20±8℃,20±10℃)和1个恒温(20℃±0℃)对照。实验结果表明:每天不同温度波动幅度对孔石莼(Ulva pertusa)的生长率具有明显的影响(P<0.05),其中(20±2)℃、(20±4)℃和(20±6)℃与恒温对照相比能明显促进孔石莼的生长,(20±10)℃则显著抑制孔石莼的生长;孔石莼相对生长率(RGR)与温度波动幅度(TA)的关系符合如下多项式方程: G=β_0+β_1(TA)+β_2(TA)~2其中G代表相对生长率,TA代表温度波动幅度,β0为方程在G轴上的截距,β_1和β2为回归系数。当平均温度为20℃时,对孔石莼生长最有利的温度波动幅度为±3.69℃。温度日节律波动对孔石莼Chl-a、Chl-b、蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05),(20±2)℃和(20±4)℃处理组的Chl-a含量显著高于恒温对照组,(20±2)℃、(20±4)和(20±6)℃处理的蛋白质和可溶性糖的绝对含量明显高于恒温对照组,但当温度波动幅度大于±8℃时,可溶性糖和细胞防御物质游离脯氨酸的含量明显增加(P<0.05)。
2.温度日节律波动对带形蜈蚣藻四分孢子放散、孢子萌发和早期发育的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。观察了带形蜈蚣藻孢子的放散,并在不同温度条件下测定了带形蜈蚣藻孢子萌发率和早期发育。实验结果表明带形蜈蚣藻四分孢子囊携四个孢子同时从藻体中排出,孢子囊膜破裂后四个孢子同时放散;四分孢子的萌发类型为Nemalion type;从带形蜈蚣藻四分孢子附着至第三次分裂,藻体的大小没有发生明显的变化;适宜的温度波动促进了带形蜈蚣藻四分孢子的萌发和早期发育。
3.温度日节律波动对真江蓠生长和营养吸收的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。研究了真江蓠在不同温度条件下的生长、元素积累和营养吸收。实验结果表明温度日节律波动能够促进了真江蓠的生长、对碳元素和氮元素的积累以及对水体中NO3--N的吸收和移除。
4.温度日节律波动中昼高温、夜低温对真江蓠营养吸收和光合作用的影响
实验共设3个温度日节律波动处理:15±3℃短光周期(L:D=8:16)、20±3℃正常光周期(L:D=12:12)、25±3℃长光周期(L:D=16:8);用15℃短光周期(L:D=8:16)、20℃正常光周期L:D=(12:12)和25℃长光周期(L:D=16:8)分别作为恒温对照。研究了在不同温度条件下昼高温和夜低温对真江蓠的营养吸收和光合作用。实验结果表明温度日节律波动中昼高温不但促进了真江蓠对水体中NO_3~--N的吸收,也促进了其净光合作用速率;同时在夜低温能够抑制真江蓠的夜间呼吸作用。上述结论能够解释温度日节律波动促进潮间带海藻生长的现象。
二、节律性干出对孔石莼生长和生化组成的影响
实验共设7个节律性干出的处理,即每12h干出0.0h、0.5h、1.0h、2.0h、3.0h、4.0h和5.0h。实验结果表明:每12h不同干出时间对孔石莼(Ulva pertusa)的生长具有明显的影响(P<0.05),其中干出0.5h、1.0h和2.0h与不干出处理相比能明显促进孔石莼的生长,干出5h则显著抑制孔石莼的生长(P<0.05);节律性干出对孔石莼叶绿素a、叶绿素b、蛋白质、可溶性糖和游离脯氨酸含量具有明显的影响(P<0.05),干出0.5h、1.0h和2.0h处理的可溶性糖的绝对含量明显高于不干出处理,干出时间过长(3.0h、4.0h和5.0h)不但使孔石莼生长物质的含量减少,并且使孔石莼的渗透溶质增加,当干出时间为5h时,可溶性糖含量突然增高;游离脯氨酸的含量也随着干出时间的增加而增加。
三、盐度日节律波动对孔石莼生长和生化组成的影响
实验共设5个盐度日节律变化处理(30±3、30±6、30±9、30±12和30±15)和一个恒盐(30±0)对照处理。实验结果表明:孔石莼能够耐受较大范围的盐度日节律性连续变化,这种变化虽然对孔石莼的生长不利,但是仍然能够使孔石莼的相对生长率保持在12%以上;每天不同盐度变化对孔石莼的生长率具有明显的影响(P<0.05),盐度变化幅度±3~±15与恒盐对照相比明显不利于孔石莼的生长,盐度变化幅度越大,孔石莼生长越缓慢。盐度日节律连续变化对孔石莼蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05)。同恒盐处理相比,盐度日节律连续变化对孔石莼叶绿素的含量的影响不明显,但叶绿素a和叶绿素b含量都有随着盐度变化幅度的增大而逐渐减少的趋势。盐度日节律性连续变化使孔石莼蛋白质和可溶性糖的相对含量显著增加(P<0.05),随着盐度变化幅度的增大,蛋白质和可溶性糖增加的趋势越明显,并且在盐度日节律性连续变化下,蛋白质增加的速度比可溶性糖增加的速度要快。同时,游离脯氨酸的相对含量随着盐度变化幅度的增加而增加,但是在缓和的盐度日节律性连续变化下(30±3)孔石莼游离脯氨酸的含量却显著小于恒盐对照组(P<0.05)。
四、光对孔石莼生长和生化组成的影响
1.光照强度对孔石莼生长和生化组成的影响
研究了50～400μmol/(m2·s)范围内光照强度对孔石莼生长和藻体生化组成的影响。研究结果表明光照强度对孔石莼的生长率具有明显的影响(P<0.05),孔石莼在光照强度为250μmol/(m~2·s)时有最大的生物量积累。光照强度低于250μmol/(m~2·s)时,孔石莼的生长率有随着光照强度的降低而减小的趋势;光照强度高于250μmol/(m~2·s)时,孔石莼的生长率随着光照强度的升高而降低(P<0.05);过高或过低的光照强度均不利于孔石莼的生长,光照强度在100μmol/(m~2·s)以下时,孔石莼的生长明显受到不利的影响,光照强度至350μmol/(m~2·s)以上、实验超过6天后藻体叶片产生溃斑,并产生了明显的光抑制。不同光照强度对孔石莼的叶绿素、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05)。随着光照强度的增加,孔石莼的叶绿素a和叶绿素b的含量逐渐变小;低光照条件可促进了孔石莼蛋白质的积累;当光照强度超过350μmol/(m~2·s)时,游离脯氨酸的含量明显增加。光照强度对孔石莼化学组成也具有明显的影响(P<0.05)。随着光照强度的升高,藻体的N元素含量有所升高,但光强达到350μmol/(m~2·s)后,随着光强的增加,N元素含量反而降低;C元素的含量在光照强度为50~350μmol/(m~2·s)范围内变化不明显,但当光照强度达到350μmol/(m~2·s)后,C元素的含量明显升高;H元素的含量在50~400μmol/(m~2·s)范围内,具有随着光照强度增加而增加的趋势。
2.光照强度日节律波动对孔石莼生长和生化组成的影响
本研究由两个实验组成。实验1共设5个光照强度波动幅度处理( 200±40μmol/(m~2·s)、200±80μmol/(m~2·s)、200±120μmol/(m~2·s)、200±160μmol/(m~2·s)和200±180μmol/(m~2·s))和1个恒定光照强度200μmol/(m~2·s)对照。实验1的结果表明:每天不同的光照强度波动幅度对孔石莼的生长率具有明显的影响( P<0.05 ),其中200±40μmol/(m~2·s)、200±80μmol/(m~2·s)和200±120μmol/(m~2·s)与恒定光照强度200μmol/(m~2·s)对照相比能明显促进孔石莼的生长;光照强度日节律波动对孔石莼Chl-a、Chl-b、蛋白质、可溶性糖和游离脯氨酸的含量具有明显的影响(P<0.05),光照波动幅度为80μmol/(m~2·s)处理下孔石莼的叶绿素a和叶绿素b的含量显著高于波动幅度为120μmol/(m~2·s)、160μmol/(m~2·s)和180μmol/(m~2·s)的处理(P<0.05),但恒定光照强度处理与光照强度波动处理下孔石莼叶绿素a和叶绿素b的含量相比没有显著的差异(P>0.05);孔石莼的蛋白质和可溶性糖含量具有随着光照强度波动幅度增大而减小的趋势。游离脯氨酸含量以恒定光照强度下最高,显著高于其余各光照强度波动处理(P<0.05)。实验2共设3个光照强度波动幅度处理(250±50μmol/(m~2·s)、250±100μmol/(m~2·s)和250±150μmol/(m~2·s))和一个恒定光照强度250μmol/(m~2·s)对照。实验2的结果表明:每天不同的光照强度波动幅度对孔石莼的生长率具有明显的影响( P<0.05 ), 250±50μmol/(m~2·s)、250±100μmol/(m~2·s)和250±150μmol/(m~2·s)处理下孔石莼终生物量和相对生长率都显著高于恒定光照强度200μmol/(m~2·s)对照组(P<0.05);恒定光照强度250μmol/(m~2·s)下孔石莼叶绿素a的含量显著小于波动幅度为50μmol/(m~2·s)和100μmol/(m~2·s)的处理(P<0.05),但与波动幅度为150μmol/(m~2·s)处理下孔石莼叶绿素a的含量相比差异不显著(P>0.05);恒定光照强度250μmol/(m~2·s)处理与光照强度波动处理下孔石莼叶绿素b的含量没有显著的差异(P>0.05)。光照波动幅度为50μmol/(m~2·s)处理下孔石莼叶绿素b的含量显著高于波动幅度为150μmol/(m~2·s)的处理(P<0.05);孔石莼的蛋白质和可溶性糖含量在恒定光照强度下都比在光照强度波动处理下高;游离脯氨酸含量以恒定光照强度下最高,显著高于其余各光照强度波动处理(P<0.05)。
3.不同光照强度下光暗周期对孔石莼生长和生化组成的影响
研究了不同光照强度下光暗周期对孔石莼(Ulva pertusa)生长和生化组成的影响。发现在长日照能够明显促进孔石莼的生长,在全光照和全黑暗处理对孔石莼的生长不利。实验1光照强度为110μmol/(m~2·s),实验2光照强度为240μmol/(m~2·s),两种光照强度下分别设置11个光暗周期处理(L:D=24:0、20:4、16:8、14:10、13:11、12:12、11:13、10:14、8:16、4:20和0:24)。实验结果表明:不同光暗周期对孔石莼的生长率具有明显的影响(P<0.05),除全光照处理外,孔石莼的生物量具有随着光照时间的增加而增加的趋势;随着日照时间的减少,叶绿素a和叶绿素b的含量都有逐渐下降的趋势;蛋白质和可溶性糖的含量在极长和极短日照长度处理中都有增加的趋势;随着光照时间的增加,游离脯氨酸的含量也逐渐增加。
4.光谱成分及其变动对孔石莼生长和生化组成的影响
研究了光谱成分及其波动对孔石莼(Ulva pertusa)生长和生化组成的影响。实验共设11个光谱成分及其波动的处理(白光、橙色光、绿色光、红色光、蓝色光、橙色光:绿色光=6:6-h、橙色光:红色光=6:6-h、绿色光:红色光=6:6-h、绿色光:蓝色光=6:6-h、红色光:蓝色光=6:6-h、蓝色光:橙色光=6:6-h)。实验结果表明:橙色光、蓝色光同白光相比对孔石莼的生长没有明显的影响,而红色光和绿色光对孔石莼的生长产生了明显的抑制,尤其以绿色光的抑制作用最为显著(P<0.05)。光谱变化处理中,绿色光:红色光=6:6-h处理出现了光增益效应。红色光和蓝色光下孔石莼的叶绿素a和叶绿素b的含量显著小于白光处理(P<0.05)。单色光的波动有利于增加孔石莼叶绿素、蛋白质和可溶性糖的含量,并能减弱游离脯氨酸在藻体内的积累。
ⅠEffects of diel fluctuating temperature on germination, growth, biochemical composition, nutrient uptake and photosynthesis of several macroalgae
1 Effects of circadian rhythms of fluctuating temperature on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under (20±2)℃, (20±4)℃, (20±6)℃, (20±8)℃and (20±10)℃circadian rhythms of fluctuating temperature conditions, and constant temperature of 20℃was used as the control. The growth rate of macroalga at (20±2)℃, (20±4)℃and (20±6)℃were significantly higher than that at constant temperature of 20℃, while growth rate at (20±8)℃and (20±10)℃were significantly lower than that at constant temperature of 20℃. The growth rate of macroalga was a quadratic function of the thermal amplitude. Such a growth model can be described by G=β_0+β_1(TA)+β_2(TA)_2 where G represents the relative growth rate, TA is thermal amplitude in degree Celsius,β0 is the intercept on the G axis, andβ_1 andβ_2 are the regression coefficients. The optimal thermal amplitude for the growth of thallus at mean temperature of 20℃was estimated to be±3.69℃. Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating temperature caused various influences (P<0.05).The content of chlorophyll, protein and total solute carbohydrate at (20±2)℃ and (20±4)℃were slightly higher than those at constant temperature of 20℃, however no statistically significant differences were found among them (P>0.05). While osmolytes (total solute carbohydrate and free proline) at (20±10)℃were significantly higher than that at 20℃(P<0.05). Therefore, more chlorophyll and carbohydrate production might account for the enhancement in the growth of macroalga at the diel fluctuating temperatures in the present study.
2 Effects of diel fluctuating temperature on germination and early growth of Grateloupia turuturu
The marcoalga Grateloupia turuturu was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The spores germination, early growth were observed in this experiment. The type of G. turutur germination belongs to Nemalion type. There was no significant difference in the spores’size between 2 cells stage and 8 cells stage after spores attached(P>0.05). The germination rate of macroalga at (15±3)℃, (20±3)℃were significantly higher than that at constant temperature of 15℃and 20℃respectively(P<0.05).
3 Effects of diel fluctuating temperature on growth and nutrient uptake of Gracilaria vermiculophylla
The marcoalga Gracilaria vermiculophylla was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The specific growth rate, nutrient uptake and NO3--N removal were determined in this experiment. The result showed that diel fluctuating temperature can enhance the growth rate, nutrient uptake and NO3--N removal of Gracilaria vermiculophylla than that under the constant temperature(P<0.05).
4 Effects of high day temperature and low night temperature on nutrient uptake and photosynthesis of Gracilaria vermiculophylla
The marcoalga Gracilaria vermiculophylla was cultured under (15±3)℃, (20±3)℃and (25±3)℃diel fluctuating temperature conditions, and constant temperatures of 15℃, 20℃and 25℃were used as the controls, respectively. The specific growth rate, nutrient uptake and photosynthetic rate and respiratory rate were determined in this experiment. The results showed that high day temperature and low night temperature play the different role during the temperature fluctuation. High day temperature not only can accelerate the NO3--N removal of Gracilaria vermiculophylla but also can enhance the net photosynthetic rate(P<0.05), and low night temperature can inhibit the respiration compare with the macroalga under the constant temperature(P<0.05).
ⅡEffects of periodical emersion on growth and biochemical composition of Ulva pertusa
The influence of emersion on the growth and biochemical composition were investigated with Ulva pertusa thalli grown under laboratory conditions with periodical emersion and submersion treatments. The accumulated biomass and daily increments of thalli with 0.5h, 1.0h and 2.0h periodical emersion every 12h were faster than those without emersion, while growth rate with 5h periodical emersion every 12h was significantly lower than that without emersion. Analysis of biochemical composition at the final stages of thalli growth revealed that periodical emersion caused various influences (P<0.05). The final content of total solute carbohydrate with 0.5h, 1.0h and 2.0h periodical emersion every 12h was significantly higher than that without emersion, while osmolyte with 5h periodical emersion every 12h was significantly higher than that without emersion (P<0.05). Therefore, mild periodical emersion could enhance the growth of Ulva pertusa, while severe periodical emersion could inhibit the growth of Ulva pertusa.
ⅢEffects of circadian rhythms of fluctuating salinity on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva Pertusa was cultured under 30±3, 30±6, 30±9, 30±12 and 30±15 circadian rhythms of fluctuating salinity conditions, and constant salinity of 30 was used as the control. The growth rate of macroalga at fluctuating salinity conditions were significantly lower than that at constant salinity of 30, while the SGR of U. pertusa at different fluctuating salinity conditions were all kept at least 12%. Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating salinity caused various influences (P<0.05). The content of chlorophyll a and chlorophyll b at fluctuating salinity conditions were slightly lower than those at constant salinity of 30, however no statistically significant differences were found among them (P>0.05). The content of protein and total solute carbohydrate at fluctuating salinity conditions were significantly higher than those at constant salinity of 30 (P<0.05), and there was a trend that the content of proline increased with the increase of amplitude of fluctuating salinity, while the content of proline at mild change of salinity (30±3) was significantly lower than that at constant salinity (P<0.05).
ⅣEffects of light on growth and biochemical composition of Ulva pertusa
1. Effects of light intensity on growth and chemical constituents of Ulva pertusa The influence of light on the growth and chemical constituents were investigated with Ulva pertusa thalli growth under laboratory conditions with different light intensity. There was a trend that the growth rate of macroalga decreased with the decrease of light intensity under 250μmol/(m~2·s), but when light intensity exceed 250μmol/(m~2·s), the growth rate of macroalga also decrased with the increase of light intensity(P<0.05). Analysis of chemical constituents at the final stages of thalli growth revealed that different light intensity caused various influences (P<0.05). There was a trend that the content of chlorophyll a and chlorophyll b decreased with the increase of light intensity. The synthesization of protein was accelerated within the lower light intensity. While osmolytes (free proline) at 350μmol/(m~2·s) and 400μmol/(m~2·s) was significantly higher than other treatments (P<0.05). The content of nitrogen increased with the increase of light intensity from 50 to 300μmol/(m~2·s), while the content of nitrogen decreased with the increase of light intensity when light intensity exceed 350μmol/(m~2·s), the content of carbon has the same trend with the trend of nitrogen, and the content of hydrogen increased with the increase of light intensity in this study.
2. Effects of circadian rhythms of fluctuating light intensity on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under 200±40μmol/(m~2·s), 200±80μmol/(m~2·s), 200±120μmol/(m~2·s), 200±160μmol/(m~2·s) and 200±180μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions, and constant light intensity of 200μmol/(m~2·s) was used as the control in experiment 1. The growth rate of macroalga at 200±40μmol/(m~2·s), 200±80μmol/(m~2·s) and 200±120μmol/(m~2·s) were significantly higher than that at constant light intensity of 200μmol/(m~2·s) (P<0.05). Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating light intensity caused various influences (P<0.05).While osmolytes (free proline) at constant light intensity of 200μmol/(m~2·s) was significantly higher than those at other treatment (P<0.05).The marcoalga Ulva pertusa was cultured under 250±50μmol/(m~2·s), 250±100μmol/(m~2·s) and 250±150μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions, and constant light intensity of 250μmol/(m~2·s) was used as the control in experiment 2. The growth rate of macroalga at 250±50μmol/(m~2·s), 250±100μmol/(m~2·s) and 250±150μmol/(m~2·s) were significantly higher than that at constant light intensity of 250μmol/(m~2·s) (P<0.05). Analysis of biochemical composition at the final stages of thaulls growth revealed that diel fluctuating light intensity caused various influences (P<0.05). .The content of chlorophyll-a at 250±50μmol/(m~2·s) and 250±100μmol/(m~2·s) was significantly higher than that at constant light intensity of 250μmol/(m~2·s) (P<0.05), the content of protein and total solute carbohydrate at 250±50μmol/(m~2·s) circadian rhythms of fluctuating light intensity conditions were significantly higher than those at constant temperature of 250μmol/(m~2·s), osmolytes (free proline) at constant light intensity of 250μmol/(m~2·s) was significantly higher than those at other treatment (P<0.05).
3. Effects of light-dark cycles on growth and biochemical composition of Ulva pertusa under different light intensity conditions
The marcoalga Ulva pertusa was cultured under L:D=24:0、20:4、16:8、14:10、13:11、12:12、11:13、10:14、8:16、4:20 and 0:24 light-dark cycles under 110μmol/(m~2·s) and 240μmol/(m~2·s) light intensity conditions, respectively. There was a trend that the growth rates of macroalga decreased with the increase of light period. The synthesization of protein and total carbohydrate were accelerated within the shorter light period and longer light period. And free proline was increased with the increase of light period.
4. Effects of light spectra and fluctuating of light spectra on growth and biochemical composition of Ulva pertusa
The marcoalga Ulva pertusa was cultured under White, Yellow, Green, Red, Blue, Yellow: Green=6:6h, Yellow: Red=6:6h, Green: Red=6:6h, Green: Blue=6:6h, Red: Blue=6:6h, Blue: Yellow=6:6h light spectra conditions. The growth rate of macroalga at Red light and Green light were significantly lower than that at White light. Emerson enhancement effect occurred at Green: Red=6:6h treatment. The content of chlorophyll, protein and total solute carbohydrate was accelerated by the fluctuating of light spectra. The content of praline was alleviated by the fluctuating of light spectra
引文
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