叶绿素荧光参数对氮、磷限制的响应及其在赤潮生消过程中的变化特征研究
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摘要
海洋中浮游植物的生长经常受到营养盐的限制。许多方法可以用来判断限制因子，但往往由于参数测定复杂，不同微藻之间存在差异等原因，主要用于室内研究。对于现场海域而言，目前主要根据水体中营养盐的浓度和比例、营养盐添加后叶绿素a（Chl a）的变化情况来判断限制因子。然而，虽然前一种方法速度较快，但是不同浮游植物对N、P的需求不尽相同，最适氮磷比也有差异，该方法只是按照一个固定的营养盐浓度和比例来判断，因此只是一个粗略的估计。而后一种方法，只是告诉我们随着浮游植物的继续生长，哪种营养元素可能会被首先耗尽，不能反映调查区域现场原位的营养状态，且一般需要几天的时间才能得出结论。因此，迫切需要建立一个快速、可靠的方法来判断现场原位营养盐的限制情况。
活体叶绿素荧光是研究光合作用的有效探针，可以快速、简便、无损伤的探测植物的光合生理状况以及各种外界因子对其产生的细微影响。其中，最常用的荧光参数为光系统Ⅱ（PSⅡ）的最大光化学量子产量（F_v/F_m）。已有研究表明，F_v/F_m对营养盐限制存在响应。但就目前来看，室内研究主要以饵料藻和淡水藻为主，近海常见藻种研究较少，且较少关注处于限制条件下的微藻重新添加营养盐后，F_v/F_m在恢复过程中的变化情况。现场海域，F_v/F_m已被认为是一个研究Fe限制的良好指标，然而应用F_v/F_m研究常量营养盐限制的报道较少，且虽有成功的案例，但结论并不统一。此外，现场海域情况复杂，除了营养盐，其他环境因子如温度、光照等同样会对F_v/F_m产生影响。而且，天然水域中同时存在多种浮游植物，群落结构也会引起F_v/F_m数值的变化。因此如何利用F_v/F_m来判断现场海域常量营养盐的限制情况，仍需进一步研究。
此外，赤潮监测过程中，浮游植物种类组成、细胞数量等参数测量必不可少，但这些参数传统的测量方法耗时较长，降低了赤潮监测的时效性。因此需要确立一个快速、可靠的指标，为赤潮的现场实时监测服务。F_v/F_m测量快速、简便。已有研究表明，赤潮生消过程中其存在变化情况。然而目前相关报道较少，仍需进一步研究。
本文采用室内培养与现场实验相结合的方法，室内选择7种我国近海常见微藻，观察了F_v/F_m对N、P限制的响应，尝试运用该参数来判断N、P限制；并将该方法应用到现场海域，判断了青岛近海的N、P限制情况。此外，以长江口邻近海域为研究区域，报道了赤潮生消过程中F_v/F_m的变化情况，尝试将该参数作为一个赤潮现场快速监测的辅助指标，为赤潮的研究提供基础资料。得到的主要结论如下：
1、室内培养条件下7种海洋微藻F_v/F_m对N、P限制的响应（1）以中肋骨条藻（Skeletonema costatum）、旋链角毛藻（Chaetoceros curvisetus）、微小原甲藻（Prorocentrum minimum）、裸甲藻（Gymnodinium sp.）、盐生杜氏藻（Dunaliella salina）、球等鞭金藻3011（Isochrysis galbana）、隐藻1533（Rhodomonas sp.）7种微藻为研究对象，分别在缺少N或P的条件下培养数天，F_v/F_m均逐渐下降。之后随着限制性营养盐的重新添加，与对照组相比，各微藻的F_v/F_m在5h~24h之内便开始显著升高（P <0.05），18h~72h之内恢复到营养盐充足时的水平；而添加其他非限制性营养盐，F_v/F_m在整个测定过程中与对照组相比没有明显差异（P>0.05）。Chl a对限制性营养盐的添加同样存在响应，但其响应时间大多晚于F_v/F_m的时间。此外，不同微藻，F_v/F_m的最大值不尽相同。
（2）根据上述结果可以推出，在浮游植物群落结构没有发生显著变化的前提下，可以根据营养盐添加后F_v/F_m的变化情况来判断现场海域是否受到N、P的限制：添加某种营养盐后，与对照组相比，如果F_v/F_m显著升高，则说明此种营养盐为限制因子；相反的，如果F_v/F_m没有显著差异，则说明此种营养盐为非限制因子。该方法速度较快，与通过营养盐加富后Chl a的变化来判断限制因子相比，更加快速。
2、青岛近海F_v/F_m对N、P限制的响应
2009年夏季在青岛近海A、B两个站位采集水样，进行了6次N、P加富实验，利用上述方法以及营养盐的浓度和比例、营养盐加富后Chl a的变化情况，三种方法来判断该区域是否受到N、P的限制。结果显示：
（1）7月24日，水体中PO4-P的浓度、F_v/F_m处于较低水平，24h后F_v/F_m、Chl a对P的添加存在响应，P为限制浮游植物生长的营养因子。8月4日，PO4-P的浓度依然处于限制水平，DIN的浓度大幅下降，B站位只有加N+P组的F_v/F_m与对照组相比显著升高，说明浮游植物的生长受到N、P的共同限制。加N+P组的Chl a浓度最高也证实了这点。9月16日各营养盐浓度有所回升，F_v/F_m数值较高。B站位F_v/F_m、Chl a对N、P的添加没有响应，说明此时N、P充足，浮游植物群落处于健康状态。A站位，F_v/F_m的初始数值、浮游植物的群落组成，与B站位类似，且添加N、P后，F_v/F_m维持在初始时的水平没有升高。说明初始时，A站位浮游植物群落也处于健康状态。随后，对照组、加N组的F_v/F_m逐渐下降，24h时显著低于加P组、N+P组的数值，说明随着浮游植物继续生长，P会被首先耗尽。此外，初始时DIN/PO4-P比值、SiO3-Si/PO4-P比值较高，48h后加P组、N+P组Chl a的浓度显著高于其他两组，也得出P为潜在限制因子的结论。
（2）在浮游植物群落结构没有发生显著变化的前提下，利用营养盐添加后F_v/F_m的变化情况来判断现场海域是否受到N、P的限制，快速（24h）、准确，还可以得到群落生理状态方面的信息。
3、长江口邻近海域春季赤潮生消过程中表层水体F_v/F_m的变化特征
2010年、2011年春季赤潮生消过程中对典型断面的表层水体进行了重复调查。研究表明，晴天时表层水体F_v/F_m存在明显的昼低夜高现象，夜晚时F_v/F_m数值最高，白天随着光强的增强F_v/F_m逐渐降低，当正午光强最强时F_v/F_m降至最低水平。暗适应1h基本上可以消除昼夜变化带来的影响，使不同时间测定的数据可以相互比较。F_v/F_m、Chl a由bbe藻类分析仪、Phyto-PAM浮游植物荧光仪两种仪器测得，研究表明两种仪器的测定结果具有较高的线性相关性。数据统一转换成Phyto-PAM的测定结果，得到的主要结果如下：
（1）2010年5月7日至5月24日期间对长江口以北的绿潮孕育区和以南的赤潮高发区进行了一次大面调查。表层水体F_v/F_m在0.19~0.74之间，整体呈现出北高南低的特点。其中，长江口以北绿潮孕育区，Chl a浓度较低，硅藻为优势种，F_v/F_m高值区分散在南北两测。长江口以南赤潮高发区，Chl a浓度较高，局部发生硅、甲藻赤潮。F_v/F_m呈现出近岸高远岸低的特点，高值区集中在沿岸的狭长地带，并在杭州湾附近海域呈舌状向外海伸展。
（2）2011年重点关注长江口以南赤潮高发区，于3月29日至5月27日硅甲藻赤潮生消期间对5个典型断面进行了9个航次的重复调查。表层水体F_v/F_m在0.17~0.73之间，F_v/F_m大于0.60的区域与赤潮区域分布类似。其中，3月29日至4月2日期间，调查海区水体普遍处于富营养化状态，高温高盐的外海水向近岸推进，从而导致外部海域水温较高，适合硅藻的生长繁殖，F_v/F_m达到0.60以上，在此区域内硅藻赤潮率先爆发。随着硅藻赤潮的发生、发展，PO4-P、SiO3-Si的浓度大幅降低，已难以满足藻细胞持续增长的需求，4月9日至4月20日期间硅藻赤潮进入消亡期，F_v/F_m普遍在0.60以下。期间，水温逐渐上升，随着长江冲淡水的补充与外扩，局部区域营养盐得到补充，F_v/F_m重新升高。5月4日至5月7日期间南部近岸区域营养盐充足，水温上升到15℃以上，适合甲藻的生长繁殖，F_v/F_m达到0.60以上，在此区域内原甲藻赤潮率先爆发。随后，5月13日至5月15日期间，F_v/F_m高值区向北移动，原甲藻赤潮向北扩张，东北部海域也发生原甲藻赤潮。5月25日至5月27日期间长江冲淡水回缩，高温高盐的外海水向近岸推进，F_v/F_m的高值区贴近岸边，在此范围内原甲藻赤潮仍在持续。
（3）温度、浊度、营养盐为影响该区域F_v/F_m的主要因素。营养盐加富实验表明，分别以骨条藻、东海原甲藻为优势的硅、甲藻群落，F_v/F_m的最大值差异不大，当F_v/F_m在0.60以上时，硅甲藻群落处于健康状态，没有受到环境因子的胁迫。否则，F_v/F_m低于0.60的水平。近岸区域主要是由于水体温度较低、浊度较高造成的；外部海域主要是由于营养盐浓度较低造成的。
（4）硅藻赤潮、原甲藻赤潮生消期间F_v/F_m表现出相似的特征：赤潮爆发前，F_v/F_m首先上升到0.60以上，然后Chl a大幅上升并爆发赤潮。赤潮期间F_v/F_m大多高于0.60的水平。赤潮消亡时，F_v/F_m重新降至0.60以下。鉴于赤潮爆发前F_v/F_m首先上升，然后Chl a大幅上升并爆发赤潮，因此可以把F_v/F_m作为藻华预警的辅助指标。
（5）硅藻在细胞氮磷比值较低的情况下更易生长，而甲藻的适应范围较宽，在细胞氮磷比值较高时，仍能达到健康状态，具有较强的生长潜力。
Growth of phytoplankton in aquatic environments is frequently limited by theavailability of nutrients. A number of methods are available for determining nutrientlimitation in phytoplankton. However, many of them are complicated, timeconsuming and vary with species. There are problems extrapolating from laboratorystudies to an understanding of nutrient limitation on natural populations.
At present, two approaches, i.e. criteria of nutrient concentrations and ratios, andnutrient enrichment bioassays using chlorophyll response as indicators, are commonlyused to assess nutrient limitation in the field. However, it is becoming increasinglyclear that the criteria of nutrient concentrations and ratios vary greatly between taxa,which introduce substantial difficulties when applying them to specific aquaticecosystems. In addition, bioassays may indicate the potential for nutrient limitation,but not necessarily show in situ nutrient status. Also, bioassays are time consumingand often require multiple days.
Chlorophyll a fluorescence allows for rapid, sensitive, and non-invasiveassessment of the physiological state of phytoplankton. The fluorescence parametermost commonly used is the maximum photochemical efficiency of photosystem Ⅱ(F_v/F_m). Changes in F_v/F_mduring nutrient limitation have been observed in numerouslaboratory cultures, but the processes involved in recovery from nutrient limitationhave been less well studied, especially the fine-scale kinetics of repair of F_v/F_mafternutrient resupply. In addition, their usefulness in the field has yet to be determined. Anumber of factors, such as light, temperature and community structure may have influences on F_v/F_m.
Here, the effects of nitrogen (N) and phosphorus (P) starvation and resupply onF_v/F_mwere examined in seven species of marine unicellular algae. F_v/F_mcoupled withnutrient enrichment experiment was used to determine nutrient limitation in thecoastal sea of Qingdao. In addition, spatial and temporal variations of F_v/F_mduringthe process of spring bloom in the adjacent area of Changjiang river estuary wereobserved. The main results are as follows:
1、 In laboratory cultures, F_v/F_mdeclined markedly under nitrate and phosphatestarvation in all species examined and recoverd completely within18h-72h ofresupply of the limiting nutrient. After24h, resupply of the limiting nutrientsignificantly increased F_v/F_mof all species above the control treatment (P <0.05),while addition of non-limiting nutrient did not change F_v/F_mrelative to the controltreatment (P>0.05). Species differences were observed in the maximal values ofF_v/F_m.
2、 F_v/F_mcoupled with nutrient enrichment experiments can be useful for rapidlydetecting N and P limitation of natural phytoplankton populations before majorchanges in community structure, and can provide information about the physiologicalstate of the community.
3、 F_v/F_mand the two above mentioned methods, i.e. nutrient criteria and bioassaysusing chlorophyll response as indicators, were used to identify limiting nutrients inthe Qingdao Olympic sailing venue. In July of2009, low DIP concentrations andF_v/F_min the initial samples coupled with F_v/F_mand Chlorophyll a (Chl a) responses toP addition indicated that P was the limiting element. In August, P was still thelimiting nutrient. In the southwestern part of the Olympic sailing venue, only the N+Ptreatment caused marked increase of F_v/F_mover the control treatment and the Chl aconcentration was highest in the N+P treatment, suggesting that N and P wereco-limiting. In September, a lack of F_v/F_mand Chl a response following nutrientenrichments together with the relatively high nutrient concentrations andphotosynthetic efficiency observed at the start of experiment suggested that N and Pwere replete and phytoplankton community was healthy in the southwestern part of the Olympic sailing venue. In the northeastern part, the initial F_v/F_mandphytoplankton community structure were similar to those in the southwestern part,and F_v/F_mdid not increase after N and P additions, indicating that phytoplanktoncommunity was in a healthy state on the day of collection. F_v/F_mand Chl a in thecontrol and N treatments were lower than those in the P and N+P treatments duringthe incubation, coupled with the high N/P ratio at the start of experiment, suggestingthat P was the potential limiting nutrient, and would be exhausted first as growthproceeded. In July of2010, neither N nor P was deficient in the southwestern part ofthe Olympic sailing venue, which was different from the result in the same period of2009.
4、The Characteristics of F_v/F_mduring the process of spring bloom in surface watersof the adjacent area of Changjiang river estuary
(1) F_v/F_mwas between0.19and0.74from7to24May2010, and ranged from0.17to0.73in the frequent blooms occurrence area, south of the Changjiang river estuaryfrom29March to27May2011. Bloom occurrence areas were mainly located onthose areas where F_v/F_mis above0.6.
(2) The primary factors affecting F_v/F_mwere temperature, turbidity and nutrients.Values of F_v/F_mgreater than0.60, were associated with healthy state of phytoplanktoncommunities. In addition, F_v/F_mchanged followed a day/night cycle with maximumvalues at night and a strong reduction at noon. Samples were dark adapted for1h atambient temperatures before measurement to avoid diel variations.
(3) The Characteristics of F_v/F_mduring the process of diatom bloom were similar tothose within the dinoflagellate bloom. F_v/F_mincreased to more than0.60before bloom.Then Chl a concentration rocketed and the bloom broke out. F_v/F_mwas above0.6inthe retention stage of bloom and decreased to below0.6in the dissipation phase. ThusF_v/F_mcan help detecting and characterizing bloom events.
(4) Dinoflagellate population can live in water mass characterized by low PO4-Pconcentration and high DIN/PO4-P ratio, but diatom population growth needs highPO4-P concentration and low DIN/PO4-P ratio. Maybe it is one of the reasons whydinoflagellate could grow better and cause bloom under lower PO4-P concentration after diatom bloom.
活体叶绿素荧光是研究光合作用的有效探针，可以快速、简便、无损伤的探测植物的光合生理状况以及各种外界因子对其产生的细微影响。其中，最常用的荧光参数为光系统Ⅱ（PSⅡ）的最大光化学量子产量（F_v/F_m）。已有研究表明，F_v/F_m对营养盐限制存在响应。但就目前来看，室内研究主要以饵料藻和淡水藻为主，近海常见藻种研究较少，且较少关注处于限制条件下的微藻重新添加营养盐后，F_v/F_m在恢复过程中的变化情况。现场海域，F_v/F_m已被认为是一个研究Fe限制的良好指标，然而应用F_v/F_m研究常量营养盐限制的报道较少，且虽有成功的案例，但结论并不统一。此外，现场海域情况复杂，除了营养盐，其他环境因子如温度、光照等同样会对F_v/F_m产生影响。而且，天然水域中同时存在多种浮游植物，群落结构也会引起F_v/F_m数值的变化。因此如何利用F_v/F_m来判断现场海域常量营养盐的限制情况，仍需进一步研究。
此外，赤潮监测过程中，浮游植物种类组成、细胞数量等参数测量必不可少，但这些参数传统的测量方法耗时较长，降低了赤潮监测的时效性。因此需要确立一个快速、可靠的指标，为赤潮的现场实时监测服务。F_v/F_m测量快速、简便。已有研究表明，赤潮生消过程中其存在变化情况。然而目前相关报道较少，仍需进一步研究。
本文采用室内培养与现场实验相结合的方法，室内选择7种我国近海常见微藻，观察了F_v/F_m对N、P限制的响应，尝试运用该参数来判断N、P限制；并将该方法应用到现场海域，判断了青岛近海的N、P限制情况。此外，以长江口邻近海域为研究区域，报道了赤潮生消过程中F_v/F_m的变化情况，尝试将该参数作为一个赤潮现场快速监测的辅助指标，为赤潮的研究提供基础资料。得到的主要结论如下：
1、室内培养条件下7种海洋微藻F_v/F_m对N、P限制的响应（1）以中肋骨条藻（Skeletonema costatum）、旋链角毛藻（Chaetoceros curvisetus）、微小原甲藻（Prorocentrum minimum）、裸甲藻（Gymnodinium sp.）、盐生杜氏藻（Dunaliella salina）、球等鞭金藻3011（Isochrysis galbana）、隐藻1533（Rhodomonas sp.）7种微藻为研究对象，分别在缺少N或P的条件下培养数天，F_v/F_m均逐渐下降。之后随着限制性营养盐的重新添加，与对照组相比，各微藻的F_v/F_m在5h~24h之内便开始显著升高（P <0.05），18h~72h之内恢复到营养盐充足时的水平；而添加其他非限制性营养盐，F_v/F_m在整个测定过程中与对照组相比没有明显差异（P>0.05）。Chl a对限制性营养盐的添加同样存在响应，但其响应时间大多晚于F_v/F_m的时间。此外，不同微藻，F_v/F_m的最大值不尽相同。
（2）根据上述结果可以推出，在浮游植物群落结构没有发生显著变化的前提下，可以根据营养盐添加后F_v/F_m的变化情况来判断现场海域是否受到N、P的限制：添加某种营养盐后，与对照组相比，如果F_v/F_m显著升高，则说明此种营养盐为限制因子；相反的，如果F_v/F_m没有显著差异，则说明此种营养盐为非限制因子。该方法速度较快，与通过营养盐加富后Chl a的变化来判断限制因子相比，更加快速。
2、青岛近海F_v/F_m对N、P限制的响应
2009年夏季在青岛近海A、B两个站位采集水样，进行了6次N、P加富实验，利用上述方法以及营养盐的浓度和比例、营养盐加富后Chl a的变化情况，三种方法来判断该区域是否受到N、P的限制。结果显示：
（1）7月24日，水体中PO4-P的浓度、F_v/F_m处于较低水平，24h后F_v/F_m、Chl a对P的添加存在响应，P为限制浮游植物生长的营养因子。8月4日，PO4-P的浓度依然处于限制水平，DIN的浓度大幅下降，B站位只有加N+P组的F_v/F_m与对照组相比显著升高，说明浮游植物的生长受到N、P的共同限制。加N+P组的Chl a浓度最高也证实了这点。9月16日各营养盐浓度有所回升，F_v/F_m数值较高。B站位F_v/F_m、Chl a对N、P的添加没有响应，说明此时N、P充足，浮游植物群落处于健康状态。A站位，F_v/F_m的初始数值、浮游植物的群落组成，与B站位类似，且添加N、P后，F_v/F_m维持在初始时的水平没有升高。说明初始时，A站位浮游植物群落也处于健康状态。随后，对照组、加N组的F_v/F_m逐渐下降，24h时显著低于加P组、N+P组的数值，说明随着浮游植物继续生长，P会被首先耗尽。此外，初始时DIN/PO4-P比值、SiO3-Si/PO4-P比值较高，48h后加P组、N+P组Chl a的浓度显著高于其他两组，也得出P为潜在限制因子的结论。
（2）在浮游植物群落结构没有发生显著变化的前提下，利用营养盐添加后F_v/F_m的变化情况来判断现场海域是否受到N、P的限制，快速（24h）、准确，还可以得到群落生理状态方面的信息。
3、长江口邻近海域春季赤潮生消过程中表层水体F_v/F_m的变化特征
2010年、2011年春季赤潮生消过程中对典型断面的表层水体进行了重复调查。研究表明，晴天时表层水体F_v/F_m存在明显的昼低夜高现象，夜晚时F_v/F_m数值最高，白天随着光强的增强F_v/F_m逐渐降低，当正午光强最强时F_v/F_m降至最低水平。暗适应1h基本上可以消除昼夜变化带来的影响，使不同时间测定的数据可以相互比较。F_v/F_m、Chl a由bbe藻类分析仪、Phyto-PAM浮游植物荧光仪两种仪器测得，研究表明两种仪器的测定结果具有较高的线性相关性。数据统一转换成Phyto-PAM的测定结果，得到的主要结果如下：
（1）2010年5月7日至5月24日期间对长江口以北的绿潮孕育区和以南的赤潮高发区进行了一次大面调查。表层水体F_v/F_m在0.19~0.74之间，整体呈现出北高南低的特点。其中，长江口以北绿潮孕育区，Chl a浓度较低，硅藻为优势种，F_v/F_m高值区分散在南北两测。长江口以南赤潮高发区，Chl a浓度较高，局部发生硅、甲藻赤潮。F_v/F_m呈现出近岸高远岸低的特点，高值区集中在沿岸的狭长地带，并在杭州湾附近海域呈舌状向外海伸展。
（2）2011年重点关注长江口以南赤潮高发区，于3月29日至5月27日硅甲藻赤潮生消期间对5个典型断面进行了9个航次的重复调查。表层水体F_v/F_m在0.17~0.73之间，F_v/F_m大于0.60的区域与赤潮区域分布类似。其中，3月29日至4月2日期间，调查海区水体普遍处于富营养化状态，高温高盐的外海水向近岸推进，从而导致外部海域水温较高，适合硅藻的生长繁殖，F_v/F_m达到0.60以上，在此区域内硅藻赤潮率先爆发。随着硅藻赤潮的发生、发展，PO4-P、SiO3-Si的浓度大幅降低，已难以满足藻细胞持续增长的需求，4月9日至4月20日期间硅藻赤潮进入消亡期，F_v/F_m普遍在0.60以下。期间，水温逐渐上升，随着长江冲淡水的补充与外扩，局部区域营养盐得到补充，F_v/F_m重新升高。5月4日至5月7日期间南部近岸区域营养盐充足，水温上升到15℃以上，适合甲藻的生长繁殖，F_v/F_m达到0.60以上，在此区域内原甲藻赤潮率先爆发。随后，5月13日至5月15日期间，F_v/F_m高值区向北移动，原甲藻赤潮向北扩张，东北部海域也发生原甲藻赤潮。5月25日至5月27日期间长江冲淡水回缩，高温高盐的外海水向近岸推进，F_v/F_m的高值区贴近岸边，在此范围内原甲藻赤潮仍在持续。
（3）温度、浊度、营养盐为影响该区域F_v/F_m的主要因素。营养盐加富实验表明，分别以骨条藻、东海原甲藻为优势的硅、甲藻群落，F_v/F_m的最大值差异不大，当F_v/F_m在0.60以上时，硅甲藻群落处于健康状态，没有受到环境因子的胁迫。否则，F_v/F_m低于0.60的水平。近岸区域主要是由于水体温度较低、浊度较高造成的；外部海域主要是由于营养盐浓度较低造成的。
（4）硅藻赤潮、原甲藻赤潮生消期间F_v/F_m表现出相似的特征：赤潮爆发前，F_v/F_m首先上升到0.60以上，然后Chl a大幅上升并爆发赤潮。赤潮期间F_v/F_m大多高于0.60的水平。赤潮消亡时，F_v/F_m重新降至0.60以下。鉴于赤潮爆发前F_v/F_m首先上升，然后Chl a大幅上升并爆发赤潮，因此可以把F_v/F_m作为藻华预警的辅助指标。
（5）硅藻在细胞氮磷比值较低的情况下更易生长，而甲藻的适应范围较宽，在细胞氮磷比值较高时，仍能达到健康状态，具有较强的生长潜力。
Growth of phytoplankton in aquatic environments is frequently limited by theavailability of nutrients. A number of methods are available for determining nutrientlimitation in phytoplankton. However, many of them are complicated, timeconsuming and vary with species. There are problems extrapolating from laboratorystudies to an understanding of nutrient limitation on natural populations.
At present, two approaches, i.e. criteria of nutrient concentrations and ratios, andnutrient enrichment bioassays using chlorophyll response as indicators, are commonlyused to assess nutrient limitation in the field. However, it is becoming increasinglyclear that the criteria of nutrient concentrations and ratios vary greatly between taxa,which introduce substantial difficulties when applying them to specific aquaticecosystems. In addition, bioassays may indicate the potential for nutrient limitation,but not necessarily show in situ nutrient status. Also, bioassays are time consumingand often require multiple days.
Chlorophyll a fluorescence allows for rapid, sensitive, and non-invasiveassessment of the physiological state of phytoplankton. The fluorescence parametermost commonly used is the maximum photochemical efficiency of photosystem Ⅱ(F_v/F_m). Changes in F_v/F_mduring nutrient limitation have been observed in numerouslaboratory cultures, but the processes involved in recovery from nutrient limitationhave been less well studied, especially the fine-scale kinetics of repair of F_v/F_mafternutrient resupply. In addition, their usefulness in the field has yet to be determined. Anumber of factors, such as light, temperature and community structure may have influences on F_v/F_m.
Here, the effects of nitrogen (N) and phosphorus (P) starvation and resupply onF_v/F_mwere examined in seven species of marine unicellular algae. F_v/F_mcoupled withnutrient enrichment experiment was used to determine nutrient limitation in thecoastal sea of Qingdao. In addition, spatial and temporal variations of F_v/F_mduringthe process of spring bloom in the adjacent area of Changjiang river estuary wereobserved. The main results are as follows:
1、 In laboratory cultures, F_v/F_mdeclined markedly under nitrate and phosphatestarvation in all species examined and recoverd completely within18h-72h ofresupply of the limiting nutrient. After24h, resupply of the limiting nutrientsignificantly increased F_v/F_mof all species above the control treatment (P <0.05),while addition of non-limiting nutrient did not change F_v/F_mrelative to the controltreatment (P>0.05). Species differences were observed in the maximal values ofF_v/F_m.
2、 F_v/F_mcoupled with nutrient enrichment experiments can be useful for rapidlydetecting N and P limitation of natural phytoplankton populations before majorchanges in community structure, and can provide information about the physiologicalstate of the community.
3、 F_v/F_mand the two above mentioned methods, i.e. nutrient criteria and bioassaysusing chlorophyll response as indicators, were used to identify limiting nutrients inthe Qingdao Olympic sailing venue. In July of2009, low DIP concentrations andF_v/F_min the initial samples coupled with F_v/F_mand Chlorophyll a (Chl a) responses toP addition indicated that P was the limiting element. In August, P was still thelimiting nutrient. In the southwestern part of the Olympic sailing venue, only the N+Ptreatment caused marked increase of F_v/F_mover the control treatment and the Chl aconcentration was highest in the N+P treatment, suggesting that N and P wereco-limiting. In September, a lack of F_v/F_mand Chl a response following nutrientenrichments together with the relatively high nutrient concentrations andphotosynthetic efficiency observed at the start of experiment suggested that N and Pwere replete and phytoplankton community was healthy in the southwestern part of the Olympic sailing venue. In the northeastern part, the initial F_v/F_mandphytoplankton community structure were similar to those in the southwestern part,and F_v/F_mdid not increase after N and P additions, indicating that phytoplanktoncommunity was in a healthy state on the day of collection. F_v/F_mand Chl a in thecontrol and N treatments were lower than those in the P and N+P treatments duringthe incubation, coupled with the high N/P ratio at the start of experiment, suggestingthat P was the potential limiting nutrient, and would be exhausted first as growthproceeded. In July of2010, neither N nor P was deficient in the southwestern part ofthe Olympic sailing venue, which was different from the result in the same period of2009.
4、The Characteristics of F_v/F_mduring the process of spring bloom in surface watersof the adjacent area of Changjiang river estuary
(1) F_v/F_mwas between0.19and0.74from7to24May2010, and ranged from0.17to0.73in the frequent blooms occurrence area, south of the Changjiang river estuaryfrom29March to27May2011. Bloom occurrence areas were mainly located onthose areas where F_v/F_mis above0.6.
(2) The primary factors affecting F_v/F_mwere temperature, turbidity and nutrients.Values of F_v/F_mgreater than0.60, were associated with healthy state of phytoplanktoncommunities. In addition, F_v/F_mchanged followed a day/night cycle with maximumvalues at night and a strong reduction at noon. Samples were dark adapted for1h atambient temperatures before measurement to avoid diel variations.
(3) The Characteristics of F_v/F_mduring the process of diatom bloom were similar tothose within the dinoflagellate bloom. F_v/F_mincreased to more than0.60before bloom.Then Chl a concentration rocketed and the bloom broke out. F_v/F_mwas above0.6inthe retention stage of bloom and decreased to below0.6in the dissipation phase. ThusF_v/F_mcan help detecting and characterizing bloom events.
(4) Dinoflagellate population can live in water mass characterized by low PO4-Pconcentration and high DIN/PO4-P ratio, but diatom population growth needs highPO4-P concentration and low DIN/PO4-P ratio. Maybe it is one of the reasons whydinoflagellate could grow better and cause bloom under lower PO4-P concentration after diatom bloom.
引文
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