春季我国不同纬度河口浮游动物群落变化趋势

根据2010-2012年春季由南向北河口海域:北仑河口、九龙江口、瓯江口、长江口和灌河口的浮游动物调查资料;通过比较不同纬度河口浮游动物群落的生物多样性指数（H'）、物种更替率（R）和群落相似系数（S）;探讨了河口浮游动物群落特征沿着不同纬度变化的趋势。结果表明;自南向北5个河口可以划分为3个不同的浮游动物区系。北仑河口和九龙江口为南部亚热带区系;位于亚热带海域;以亚热带种为主;H'分别为1.01和2.18;两河口之间R值为65.52%;S值为51.28%;瓯江口和长江口为中部过渡区系;位于亚热带和温带过渡海域;亚热带种和暖温带种是主要出现的种类;H'分别为1.31和1.86;两河口之间R值为58.33%;S值为58.82%;灌河口单独为北部温带区系;位于温带海域;暖温带种是主要出现的种类;H'值为1.19;与其余河口物种更替率均高于80.00%;群落相似系数均低于32.00%;表明群落结构由南到北的变化非常显著。线性回归结果表明:我国不同河口之间浮游动物的物种更替率与纬度差（两河口所处纬度的差值;δN）呈正相关关系（R=0.026ΔN+0.608;p=0.002）;群落相似系数与纬度差呈负相关关系（S=0.034ΔN+0.578;p=0.001）。春季;我国不同纬度河口浮游动物群落结构的变化;主要是因为影响各河口的水温和水团存在较大差别所致。     

2014年春季黄、渤海浮游动物群落特征

为了解黄、渤海浮游动物群落特征, 利用2014年春季在黄、渤海采集的浮游动物样品, 对浮游动物的种类组成、丰度、生物量、优势种和生物多样性进行分析, 分析浮游动物群落结构与环境因子间的相关关系, 并划分了浮游动物群落。共记录浮游动物78个种类(南黄海69种、北黄海42种、渤海47种), 浮游幼虫占比最高。研究海域的浮游动物平均丰度为3 183.8±7 829.7 ind./m³, 剔除夜光虫(Noctiluca scintillans)后, 平均丰度为537.3±554.2 ind./m³; 平均生物量为409.1±517.5 mg/m³。夜光虫、中华哲水蚤(Calanus sinicus)和强壮箭虫(Sagitta crassa)是黄、渤海的共同优势种。结合香农-威纳指数和累积优势度曲线分析, 渤海浮游动物的生物多样性最低, 北黄海次之, 南黄海的多样性最高。聚类分析表明, 研究海域浮游动物群落可划分为渤-黄海沿岸群落、黄海中部群落和南黄海-东海近岸混合水群落。相关性分析显示, 研究海域与浮游动物群落结构相关性最高的环境因子组合为温度、盐度和叶绿素a浓度。     

基于延绳钓观察员数据的热带太平洋中东部食物网拓扑结构分析

热带太平洋是金枪鱼类、旗鱼类等渔业的主要作业海域，但目前针对此海域食物网的研究仍十分缺乏。本文根据2017年6月-11月中国金枪鱼科学观察员在热带太平洋中东部海域（13°09''N -34°27''S, 100°03''-176°17''W）采集的鱼类样品，结合胃含物及其他分析方法以35个物种间的摄食关系为基础对此海域的食物网结构进行了分析。拓扑学指标（节点强度，D；中心性指数，BC和CC；拓扑重要性指数，TI¹和TI³；关键性指数，K、K_t和K_b）和Key-Player算法（KPP-1和KPP-2）用来筛选关键种，并结合体长数据建立了基于关键种的简化食物网。肯德尔相关性分析和聚类分析结果表明，不同拓扑学指标筛选出的关键种是一致的。大多数关键种与太平洋其他海域的研究结果是相同的，如剑鱼（Xiphias gladius）、长鳍金枪鱼（Thunnus alalunga）、小吻四鳍旗鱼（Tetrapturus angustirostris）头足类和鲭属。移除5个关键种后，食物网将被离散为许多互不连接的部分（F=0.632，F^D=0.795，R^D=0.957），表明这些物种在维持食物网结构和稳定性中的必要性。物种体长是本研究构建简化食物网时的决定性因素。研究结果有助于提高对热带太平洋食物网结构的认识，同时为进一步的生态系统动力学研究提供科学依据。     

2011—2020年调水调沙前后黄河口海域浮游动物年间变化及影响因子

依据2011—2020年(2016年中断)黄河调水调沙工程实施前、后黄河口海域18个站位野外调查数据, 研究浮游动物种类组成和丰度变化及优势种更替, 以查明黄河口海域浮游动物的年间变动规律及调水调沙工程的影响。结果显示: 共记录浮游动物82种类(包括浮游幼体16类), 其中刺胞动物种类数最多, 共记录26种, 其次是桡足类出现23种。浮游动物种类组成的年间变化差异显著(P < 0.05), 2017年(未进行调水调沙)最低; 调水调沙前、后分别记录浮游动物69种和75种, 调水调沙前后种类数呈正相关(r=0.684, P=0.042); 调水调沙前后优势种更替率较高; 浮游动物丰度在2011—2015年较高, 2017—2020年相对较低, 调水调沙后浮游动物丰度年间变化达极显著水平(P < 0.01), 而调水调沙前后没有显著差异; 浮游动物丰度水平分布总体呈现从黄河口近岸到离岸递减趋势, 密集区位于黄河口北部近岸海域, 调水调沙后密集中心较调水调沙前向离岸海域略有推移。冗余分析表明, 对黄河口浮游动物年间变化影响较大的因子是叶绿素a浓度、溶解氧浓度和水温。研究表明: 2011—2020年调水调沙期间黄河口海域浮游动物丰度年间变动大于调水调沙前后变动, 调水调沙实施对浮游动物影响持续时间较短。     

AN ANALYSIS OF THE BENTHIC FAUNAL COMMUNITY STRUCTURE IN SOFT-BOTTOM INTERTIDAL ZONE ALONG NORTHERN COAST OF HANGZHOU BAY

Four indexes of species diversity (H''), species richness (d), species evenness (J) and species dominance (D₂) are used in this paper to analyse the.characteristics of the benthic faunal community structure in the soft-bottom intertidal zone along the northern coast of the Hang-zhou Bay.The results show a rather simple benthic faunal community structure in the survey area, with the average values of the four indexes from the five transects, namly, H''=2.06, d=3.34, J=0.46 and D₂=0.73.Different degrees of the seasonal and regional variations have been observed from each index per transect.     

瓯江河口盐度层化数值研究

瓯江口是一个径流量变化剧烈的强潮河口。本文基于非结构网格FVCOM模型，建立瓯江口海域大范围三维数学模型，研究不同时间尺度（潮周期、大-小潮）的盐度变化，并利用势能异常动力方程对数值模拟结果分析了瓯江口层化过程的动力机制。同时，利用河口R_i数和层化参数△s/<s>研究了不同时间尺度的层化稳定性及其空间变化，得出决定层化状态的潮差和径流量的阈值。结果显示：瓯江北口上段、中段和口门在潮差分别超过3.8m、4.0m和4.6m时呈完全混合状态。当径流量小于280 m³/s或大于510 m³/s，北口上段持续完全混合；而在口门附近，完全混合和层化的临界径流量约为280 m³/s。研究认为瓯江河口北口存在周期性的层化，北口下段在落潮和涨潮初期呈部分混合状态，而其它时段为完全混合。上段只在落潮初期存在层化。层化增强主要是纵向对流与横向速度剪切导致，而湍混合和纵向潮应力是层化减弱的主要因素。     

夏季闽江口和椒江口浮游动物群落结构的比较

根据2008年9月在闽江口水域和2010年8月在椒江口水域的调查资料,对两个河口浮游动物的种类组成、生态类群、优势种和生态适应差异进行分析比较,探讨夏季闽江口海域和椒江口海域浮游动物群落结构的区别及其组成受水团变化的影响。研究结果表明:夏季,椒江口受上升流的影响出现13个暖温带种,导致椒江口(44)水域浮游动物种类数要大于闽江口(36);闽江口海域夏季主要受到台湾暖流的影响,6个优势种全部为亚热带种,椒江口有7个优势种,包括4个亚热带种和3个暖温带种;从适温性上看,闽江口和椒江口均主要以亚热带种为主,椒江口由于上升流的原因出现较多数量的暖温带种,从适盐性上看,椒江口主要以近海种为主,闽江口主要以外海种为主,这是闽江口和椒江口浮游动物生态类群的主要差异;闽江口浮游动物群落结构主要受到台湾暖流和沿岸流的影响,而椒江口主要受到浙江近海上升流和沿岸流的影响,通过对两个河口群落结构的比较和生态适应差异的分析,表明水团是各自浮游动物群落结构的形成过程中的一个重要因素。     

New Relationship for Estimation of Wave Overtopping on Vertical Walls

Soft computing tools in the form of combination of multiple nonlinear regression and M5'' model tree were used for estimation of overtopping rate at the vertical coastal structures. For reliable and precise estimation of overtopping rate, the experimental data available in the database CLASH were used. The dimensionless overtopping rate was estimated in terms of conventional dimensionless parameters including the relative crest freeboard R_c/H_s, seabed slope tanθ, deep water wave steepness S_om, surf similarity ξ_om and local relative water depth h_t/H_s. The accuracy of the new model was compared with other existing models and also evaluated with some field measurements. The results indicated that the model presented in this paper is more accurate than other existing models. With statistical parameters, it is shown that the accuracy of predictions in the new model is better than that of other models.     

2009 年春季辽东湾网采浮游植物群落结构

分析了2009 年春季(5 月)渤海辽东湾网采浮游植物物种组成, 丰度分布, 生物多样性指数及群落相似性。春季共鉴定出浮游植物3 门23 属32 种, 主要隶属硅藻门和甲藻门, 生态类型多为温带近岸型和广布型种类, 少部分为暖海性物种或大洋性物种。优势种有夜光藻(Noctiluca scientillans)、翼鼻状藻(Proboscia alata)、旋链角毛藻(Chaetoceros curvisetus)、巨圆筛藻(Coscinodiscus gigas)和尖刺伪菱形藻(Pseudo-nitzschia pungens)等。浮游植物在各站位的细胞丰度分布不均匀, 其平面分布呈现由沿岸海域向海湾中部海域递减的趋势。Shannon-Wiener 指数、Pielou 指数较低, 群落结构简单, 稳定性较差。聚类分析结果表明群落之间的相似性较高。     

青岛董家口港邻近海域春秋季浮游动物群落结构特征

以2020年11月（秋季）和2021年3月（春季）在青岛董家口港邻近海域进行的浮游动物调查数据为基础,分析了该海域浮游动物的群落结构特征。研究海域春、秋两季总共发现浮游动物33种成体,11类浮游幼虫,其中春季调查发现18种成体,5类浮游幼虫,秋季发现24种成体,9类浮游幼虫;春、秋两季调查浮游动物平均个体密度分别为129.4、69.4 ind/m³;平均生物量（湿重）为1 166.5、23.8 mg/m³;平均种类多样性指数（H′）为1.38、2.61;丰度（d）为0.95、1.77;均匀度（J）为0.90、0.67;春季调查优势种为八斑芮氏水母（Rathkea octopunctata）、中华哲水蚤（Calanus sinicus）、腹针胸刺水蚤（Centropages abdominalis）、强壮滨箭虫（Aidanosagitta crassa）,秋季调查优势种为五角水母（Muggiaea atlantica）、双生水母（Diphyes chamissonis）、拟长腹剑水蚤（Oithona similis）、小拟哲水蚤（Paracalanus parvus）、异体住囊虫（Oikopleura dioica）、近缘大眼剑水蚤（Corycaeus affinis）、多毛类幼体、蛇尾纲长腕幼虫;聚类分析表明调查海域浮游动物可划分为两个组。     

Phylogenetic and morphometric relationships between two species of genus Auxis from the South China Sea and Java Sea

This study was aimed at examining the phylogenetic and morphometric relationship of frigate tuna(Auxis thazard) and bullet tuna(Auxis rochei) using mt DNA D-loop(control) region sequences and truss measurements,respectively. Maximum-likelihood(ML) tree and median-joining network showed that haplotypes from Auxis populations grouped separately. Discriminant function analyses and non-metric multidimensional scaling of morphometric data showed that bullet and frigate tuna were differed mainly with respect to truss measurements on the anterior region, suggesting adaptation to different diets available in different environments. Historic demographic analyses of sequence data showed that both bullet and frigate tuna had not undergone a significant sudden population expansion recently. Non-significant value of Tajimas' s D and Fu's F_S were indicated an effective large and stable population size for longer period of both species in South China Sea and Java Sea species. This study gives first report on the complementary relationship between morphometric and genetic analysis in stock discrimination of genus Auxis.     

南海北部亚热带滨珊瑚对低温的耐受能力

Marginal scleractinian corals growing at their latitudinal limits should be quite sensitive to variations in winter sea surface temperatures(SSTs). An extreme cold event occurring in early 2008 offered a unique opportunity to examine the effect of cold-water anomalies on Porites lutea corals and their physiological tolerance and acclimation in the subtropical northern South China Sea(NSCS). Besides in-situ observation, a subsequent aquarium-based experiment was designed for reproducing the chilling process and a 50-year-long Sr/Ca ratio profile from two P. lutea skeletal slabs was analyzed for reconstructed the historical annual minimum SSTs which ceased Porites calcification. The 2008 low-temperature anomaly caused the minimum daily mean SSTs dropped below 13°C in the Daya Bay. The stress symptoms displayed by local P. lutea colonies included polyp retraction, reduced coloration and pale, but none showed tissue sloughing. The ability of P. lutea to survive implied its tolerance of extreme low temperatures. Here we suggest a model on the tolerance of high-latitude Porites under low-temperature stresses, which is when SSTs drop below 18°C, Porites corals contract their tentacles(losing heterotrophic capability), then cease calcification(reducing energy consumption), and meanwhile maintain relatively high levels of zooxanthellae density(sustaining host's life via photosynthetic capacity of symbiotic zooxanthellae). This study revealed remarkable acclimatization of P. lutea corals to low temperature extremes. This acclimatization is beneficial for Porites corals in the NSCS to expand their living ranges towards the higher-latitude areas and have the potential to be the incipient reef former.     

东海春季赤潮过程中优势种的演替:生物-物理耦合数值模拟

In the East China Sea(ECS), the succession of causative species responsible for blooms is a recurrent phenomenon during the spring, which changes from diatoms to dinoflagellates. Observations from space and in situ cruises captured this pattern of succession during spring of 2005. In this study, we coupled two biological models, which were developed previously for Skeletonema costatum and Prorocentrum donghaiense,into a circulation model tailored for the ECS. The coupled biophysical model was used to hindcast the blooms and to test the hypothesis proposed in earlier studies that phosphate(PO4 3–) is the first-order decider of the succession. The coupled model successfully reproduced the hydrodynamics(as described in a companion paper by Sun et al.(1), the spatiotemporal distribution of the chlorophyll a(Chl a) concentration, and the species succession reasonably well. By analyzing the effects of different factors on the surface Chl a distribution, we confirmed that the offshore boundaries of the blooms were confined by PO4 3–. In addition, we suggest that surface wind fields may modulate the horizontal distribution of blooms. Thus, during the dispersal of blooms, surface winds coupled with PO4 3– may control the succession of blooms in the ECS. The proposed coupled model provides a benchmark to facilitate future improvements by including more size classes for organisms, multiple nutrient schemes, and additional processes.     

温度、盐度和光照对浒苔生长和光合作用的影响

Intensive Pyropia aquaculture in the coast of southwestern Yellow Sea and its subsequent waste, including disposed Ulva prolifera, was speculated to be one of the major sources for the large-scale green tide proceeding in the Yellow Sea since 2007. It was, however, unclear how the detached U. prolifera responded and resumed growing after they detached from its original habitat. In this study, we investigated the growth and photosynthetic response of the detached U. prolifera to various temperature, salinity and irradiance in the laboratory. The photosynthetic rate of the detached U. prolifera was significantly higher at moderate temperature levels(14–27°C)and high salinity(26–32), with optimum at 23°C and 32. Both low(14°C) and highest temperature(40°C), as well as low salinity(8) had adverse effects on the photosynthesis. Compared with the other Ulva species, U. prolifera showed higher saturated irradiance and no significant photoinhibition at high irradiance, indicating the great tolerance of U. prolifera to the high irradiance. The dense branch and complex structure of floating mats could help protect the thalli and reduce photoinhibition in field. Furthermore, temperature exerted a stronger influence on the growth rate of the detached U. prolifera compared to salinity. Overall, the high growth rate of this detached U. prolifera(10.6%–16.7% d~(–1)) at a wide range of temperature(5–32°C) and salinity(14–32) implied its blooming tendency with fluctuated salinity and temperature during floating. The environmental parameters in the southwestern Yellow Sea at the beginning of green tide were coincident with the optimal conditions for the detached U. prolifera.     

“海参-对虾”混养海水池塘的食物网结构及营养层级研究

The food sources of aquacultured Apostichopus japonicus and the trophic levels of organisms in a sea cucumber(A. japonicus) and prawn(Penaeus japonica) polyculture system in a saltwater pond in Zhuanghe, Liaoning Province were examined using carbon and nitrogen stable isotopes. Across organisms, δ13C ranged from(–25.47±0.20)‰ to(–16.48±0.17)‰(mean±SD), and δ15N ranged from(4.23±0.49)‰ to(12.44±0.09)‰. The δ13C and δ15N contents of A. japonicus, P. japonica and Fenneropenaeus chinensis were comparatively higher than those of other organisms. Values of δ13C and δ15N revealed that P. japonica, Hemigrapsus sanguineus and Neomysis japonica comprised the largest component of the diet of A. japonicus. The mean trophic level of the organisms in this saltwater pond polyculture system was(2.75±0.08). P. japonica, A. japonicus, F. chinensis,Synechogobius hasta and Neomysis japonica were in the 3rd trophic level(2–3); jellyfish, H. sanguineus and zooplankton were in the 2nd trophic level(1–2); and Enteromorpha prolifera, benthic microalgae, periphyton and suspended matter primarily consisting of phytoplankton, bacteria and humus were in the primary trophic level(0–1).     

大型水母沙海蜇暴发的温带贫营养海洋生态系统中细菌和浮游植物的关系

Bacterial abundance, phytoplankton community structure and environmental parameters were investigated to study the relationships between bacteria and phytoplankton during giant jellyfish Nemopilema nomurai blooms in the central Yellow Sea during 2013. N. nomurai appeared in June, increased in August, reached a peak and began to degrade in September 2013. Results showed that phosphate was possible a key nutrient for both phytoplankton and bacteria in June, but it changed to nitrate in August and September. Phytoplankton composition significantly changed that pico-phytoplankton relative biomass significantly increased, whereas other size phytoplankton significantly decreased during jellyfish bloom. In June, a significantly positive correlation was observed between chlorophyll a concentration and bacterial abundance(r=0.67, P0.001, n=34).During jellyfish outbreak in August, there was no significant correlation between phytoplankton and bacteria(r=0.11, P0.05, n=25), but the relationship(r=0.71, P0.001, n=31) was rebuilt with jellyfish degradation in September. In August, small size phytoplankton occupied the mixed layer in offshore stations, while bacteria almost distributed evenly in vertical. Chlorophyll a concentration significantly increased from(0.42±0.056) μg/L in June to(0.74±0.174) μg/L in August, while bacterial abundance just slightly increased. Additionally, the negative net community production indicated that community respiration was not entirely determined by the local primary productivity in August. These results indicated that jellyfish blooms potentially affect coupling of phytoplankton and bacteria in marine ecosystems.     

南极西北威德尔海上层海洋湍流混合研究

Turbulent mixing in the upper ocean(30-200 m) of the northwestern Weddell Sea is investigated based on profiles of temperature,salinity and microstructure data obtained during February 2014.Vertical thermohaline structures are distinct due to geographic features and sea ice distribution,resulting in that turbulent dissipation rates(ε) and turbulent diffusivity(K) are vertically and spatially non-uniform.On the shelf north of Antarctic Peninsula and Philip Ridge,with a relatively homogeneous vertical structure of temperature and salinity through the entire water column in the upper 200 m,both ε and K show significantly enhanced values in the order of O(10~(-7))-O(10~(-6)) W/kg and O(10~(-3))-O(10~(-2)) m~2/s respectively,about two or three orders of magnitude higher than those in the open ocean.Mixing intensities tend to be mild due to strong stratification in the Powell Basin and South Orkney Plateau,where s decreases with depth from O(10~(-8)) to O(10~(-9)) W/kg,while K changes vertically in an inverse direction relative to s from O(10~(-6)) to O(10~(-5)) m~2/s.In the marginal ice zone,K is vertically stable with the order of10~(-4) m~2/s although both intense dissipation and strong stratification occur at depth of 50-100 m below a cold freshened mixed layer.Though previous studies indentify wind work and tides as the primary energy sources for turbulent mixing in coastal regions,our results indicate weak relationship between K and wind stress or tidal kinetic energy.Instead,intensified mixing occurs with large bottom roughness,demonstrating that only when internal waves generated by wind and tide impinge on steep topography can the energy dissipate to support mixing.In addition,geostrophic current flowing out of the Weddell Sea through the gap west of Philip Passage is another energy source contributing to the local intense mixing.     

Biology and distribution of hyperiids in the Sea of Okhotsk

Long-term studies in the Sea of Okhotsk (1986–2012) demonstrated that hyperiids account for a small portion of zooplankton, making up only from 1.0 to 5.3% by weight. The minimum hyperiid biomass was observed in the coastal zone. The hyperiid biomass in the open water shelf community increased from spring to autumn. In the Sea of Okhotsk, hyperiids are represented by ten species, among which the most abundant in the northern regions (mainly in Shelikhov Gulf) is Themisto libellula, while the most abundant in other regions is T. pacifica. The distribution of T. pacifica in different seasons showed that the maximum biomass was concentrated in the deep-water zone. The range of T. libellula in the Sea of Okhotsk is mostly limited to the Shelikhov Gulf, but in certain years its habitation area can expand. In warm years with low ice coverage, the maximum expansion of T. libellula to the west in the shelf zone of the northern part of the Sea of Okhotsk was observed. Since variability of salinity and temperature in the northern part of the Sea of Okhotsk may be fatal for the cryophilic stenohaline T. libellula species, forecasted changes in thermohaline circulation will make it possible to predict the population dynamics of this important species.     

Prokaryotic community composition revealed by denaturing gradient gel electrophoresis in the East Sea

To understand the temporal and spatial variation of the prokaryotic community in the East Sea, their composition was determined by polymerase chain reaction (PCR)-denaturing gradient gel electrophoresis (DGGE)-sequencing techniques. The investigations were conducted twice annually in 2007 and 2009 in coastal and offshore stations. Prokaryotic abundance (PA), leucine incorporation rate, and other environmental parameters were also measured. By using the DGGE approach, we obtained 283 bacterial sequences and 160 archaeal sequences. The most frequently detected bacterial phylotypes during the investigations belonged to Alphaproteobacteria, Gammaproteobacteria, and Bacteroidetes. However, their relative compositions differed in time and space. Although Alphaproteobacteria and Bacteroidetes were the dominant groups in the surface water in May 2007 and in May and October 2007, Gammaproteobacteria was dominant in mesopelagic samples. However, Gammaproteobacteria was overwhelmingly dominant in most samples in August 2009. Although Deltaproteobacteria was rarely found as a dominant bacterial group, it occupied the highest fraction in a mesopelagic sample in October 2007. Epsilonproteobacteria also showed a similar trend, although its maximal dominance was found in a mesopelagic sample in August 2009. The archaeal community was dominated overwhelmingly by members of the Euryarchaeota in most of the investigations. However, Nitrosopumilales was dominant in aphotic samples in August 2009. Further, their spatiotemporal composition at the family level changed more dynamically in the East Sea. These temporal and spatial distributions of the prokaryotic community were influenced mainly by seawater temperature and depth in the East Sea.     

Measuring chlorophyll <Emphasis Type="Italic">a</Emphasis> concentrations in the Sea of Japan using probe and flow fluorimeters

The spatial variability of chlorophyll a concentrations was studied from the data of two near-shore expeditions and the cruise of the R/V Akademik M.A. Lavrent’ev in October–November 2010 over the northwestern part of the Sea of Japan. The sections across eddies showed a maximum of chlorophyll a at a depth of 40 m. According to the data from the cruise, the chlorophyll a concentration was maximum in the north of the sea and decreased to the south. In parallel, the procedures for chlorophyll a determination were compared for spectrophotometry with a fluorescence probe and a fluorescence flow system. The probe data of chlorophyll a fluorescence showed a high correlation with the chlorophyll a concentrations by spectrophotometry. On the contrary, data on chlorophyll a concentrations from spectrophotometry did not agree with those from the flow system. It was shown that a fluorimeter in the flow system recorded dissolved organic matter along with the chlorophyll a fluorescence.