颗石藻颗石粒形态的原子力显微观测方法: 以赫氏艾密里藻为例

颗石藻(coccolithophore)作为一种模式生物, 在重建古海洋气候和环境以及预测未来全球气候变化中起着很重要的作用, 赫氏艾密里藻(Emiliania huxleyi)是颗石藻最为典型的代表种。钙质颗石粒(coccolith)是颗石藻形态分类的主要依据, 有着非常精细和复杂的结构, 在样品收集过程中很容易遭到破坏, 这是颗石藻鉴定中经常遇到的一个技术问题。国际上还没有统一的颗石藻定量采样和样品分析方法。本文采用原子力显微方法(atomic force microscopy, AFM)对赫氏艾密里藻的颗石粒形态进行了超显微观察研究, 获取不同扫描范围的高度图(height image)和形貌图(deflection image)以观测其形态结构, 并建立了针对颗石藻的原子力显微样品制备方法。通过离心与膜过滤两种方法收集赫氏艾密里藻, 比较后得出了一种简单、快速的适合于观测颗石藻在大气环境成像的样品处理、制备和图像采集方法: 3,000-4,000 rpm, 20℃离心5 min, 收集颗石藻, 去除有机杂质后取白色沉淀, 将沉淀物悬浮于0.05 M NH₄HCO₃溶液中, 悬浮液滴加于盖玻片表面, 20℃晾干后于样品台在AFM接触模式(contact mode)下原子级扫描, 扫描范围50 µm, 频率1 Hz, 可以得到优质的颗石粒形态图像, 有助于颗石藻的分类鉴别。该方法可用于室内不同环境梯度或参数下的颗石粒形态结构及颗石藻藻华的检测与研究。     

中国近海今生颗石藻物种多样性初步研究

本文统计了中国近海97种今生颗石藻的记录,分别隶属于4个目,11个科,44个属.其中条结藻属(Syracosphaera)包含物种数最多,达20种.本文记录的中国近海5个今生颗石藻新记录种是:莱卡曲球藻(Cyrtosphaera lecaliae)、戏剧条结藻(Syracosphaera histrica)、突筋科球藻...     

2006年秋季东海陆架浮游植物群集

根据2006年11月19日—12月24日在东海陆架海域（24.0°—32.0°N,120.0°—127.0 °E）进行的综合采样调查,对调查海域浮游植物的群集特征进行了研究.本次调查共发现浮游植物4门64属145种（包括未定名种）,其中硅藻是该调查海区浮游植物的主要功能群,其次为甲藻.主要的优势种为菱形海线藻、圆海链藻、丹麦细柱藻、斯氏几内亚藻、尖刺伪菱形藻和铁氏束毛藻等.浮游植物细胞丰度为0.09～5.11 cells·ml^-1,平均值为4.92 cells·ml^-1,秋季东海陆架浮游植物细胞丰度的平面分布主要由浮游硅藻刻画,其高值出现在调查区的北、东部离岸海域,细胞丰度大于30 cells·ml^-1.浮游植物及硅藻和甲藻的细胞丰度垂直变化趋势基本上都是随着水深增加,丰度逐渐减小.各个断面细胞丰度密集区主要集中在表层,个别站位的中层和底层也出现浮游植物密集区.调查区浮游植物的多样性指数平均值为2.80,其高值位于调查区东北和东南部,物种均匀度指数平均值为0.81,其高值位于调查区中部,生物多样性的分布与细胞丰度的分布一致.从Pearson相关性指数来看,亚硝酸盐、溶解态有机氮(DON)和温度显著影响调查区浮游植物的生长.     

颗石球缓解紫外辐射对颗石藻光合作用的胁迫

正颗石藻(Coccolithophore)是一类胞外具有数层钙质颗石粒(Coccolith)的海洋钙化金藻,在绝大部分海域均有分布。颗石藻通过光合固碳向深海沉降有机碳颗粒,同时其钙化作用形成的颗石粒是海底的重要沉积物。它们与钙化浮游动物翼足类(Petropod)钙化量可占海洋生物钙化量的80%,对于海洋碳、钙循环有重要意义~([1,2])。颗石藻在条件合适时,可形成大面积藻华(全球每年可达上百万平方公里)~([3])。促进藻华形成的环境因素很     

2006年秋季东海陆架浮游植物群集

根据2006年11月19日--12月24日在东海陆架海域(24.0°--32.0°N,120.0°--127.0°E)进行的综合采样调查,对调查海域浮游植物的群集特征进行了研究.本次调查共发现浮游植物4门64属145种(包括未定名种),其中硅藻是该调查海区浮游植物的主要功能群,其次为甲藻.主要的优势种为菱形海线藻、圆海链藻、丹麦细柱藻、斯氏几内亚藻、尖刺伪菱形藻和铁氏束毛藻等.浮游植物细胞丰度为0.09～5.11 cells·ml-1,平均值为4.92 cells·ml-1,秋季东海陆架浮游植物细胞丰度的平面分布主要由浮游硅藻刻画,其高值出现在调查区的北、东部离岸海域,细胞丰度大于30 cells·ml-1.浮游植物及硅藻和甲藻的细胞丰度垂直变化趋势基本上都是随着水深增加,丰度逐渐减小.各个断面细胞丰度密集区主要集中在表层,个别站位的中层和底层也出现浮游植物密集区.调查区浮游植物的多样性指数平均值为2.80,其高值位于调查区东北和东南部,物种均匀度指数平均值为0.81,其高值位于调查区中部,生物多样性的分布与细胞丰度的分布一致.从Pearson相关性指数来看,亚硝酸盐、溶解态有机氮(DON)和温度显著影响调查区浮游植物的生长.     

海洋中的精灵——颗石藻

当恐龙称霸的时期,在海洋中生活着一类奇特的微体生物——颗石藻。颗石藻是海洋中重要的单细胞浮游植物类群,通常具有碳酸钙的外壳（称之为颗石粒,下文有详解）,藻体整体多为椭球形,少数物种呈现梭形等其他形态,大小多介于2～20um,     

2009年冬季东海浮游植物群集

2009年12月23日—2010年1月5日在东海海域(24°00’—32°00’N,120°00’—128°00’E)68个站位进行了水文、化学和生物的综合调查,应用Utermhl方法对调查海域的浮游植物群集进行了研究。经284个浮游植物采水样品的分析,共发现浮游植物4门67属171种(含13个未定种)。浮游植物群集主要由硅藻和甲藻组成,还有少量的蓝藻和金藻,物种以沿岸广温型为主,优势种为:具槽帕拉藻(Paralia sulcata)、菱形海线藻(Thalassionema nitzschioides)、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)、安哥拉海链藻(Thalassiosira angulata=并基海链藻Thalassiosira decipens)和细弱海链藻(Thalassiosira subtilis)等。调查浮游植物细胞丰度介于0.356×103—142.578×103个/L,平均值为14.137×103个/L;硅藻占浮游植物细胞丰度的比例最大,介于0.356×103—142.578×103个/L,平均值为13.023×103个/L;其次为甲藻,细胞丰度介于0.356×103—11.378×103个/L,平均值为1.177×103个/L。调查海域表层浮游植物细胞丰度的平面分布由硅藻刻画,高值区出现在调查区北部即长江口东北部海域,甲藻在调查区南部和东南部丰度较高。细胞丰度在水体中的垂直分布趋势为先上升后下降,最大值出现在10m层。从断面分布上看,细胞丰度在调查区近岸和远岸较高、中部较低。Pearson相关性分析表明,调查区浮游植物细胞丰度与磷酸盐和硅酸盐浓度呈显著正相关,与温度呈显著负相关,与硝酸盐相关性不明显。     

海洋中的精灵海灵——颗石藻

正当恐龙称霸的时期,在海洋中生活着一类奇特的微体生物——颗石藻。颗石藻是海洋中重要的单细胞浮游植物类群,通常具有碳酸钙的外壳(称之为颗石粒,下文有详解),藻体整体多为椭球形,少数物种呈现梭形等其他形态,大小多介于2～20μm,少数大于20μm;但碳酸钙外壳具有多种形态,这也是形态学鉴定的主要依据。现已知最早的颗石藻化石大约出现在距今2.2亿年左右,自那时起,颗石藻适应了当时的海洋环境,通过进化产生了由体内向体外     

夏、冬季南海北部浮游植物群落特征

对2009年7月19日—8月16日和2010年1月6—30日南海北部(18°—23.5°N、109°—120°E)两个航次的浮游植物样品应用Utermhl方法进行了分析鉴定。结果如下:夏季样品鉴定浮游植物4门72属150种,浮游植物细胞丰度范围为(0.16—6001.78)×103个/L,平均细胞丰度为26.49×103个/L,硅藻的平均细胞丰度为25.81×103个/L,主要优势种属有铁氏束毛藻(Trichodesmium thiebautii)、菱形海线藻(Thalassionema nitzschioides)、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)及裸甲藻(Gymnodinium spp.)等;冬季样品鉴定浮游植物4门58属168种,浮游植物细胞丰度范围为(0.08—37.52)×103个/L,浮游植物平均细胞丰度为2.69×103个/L,硅藻的平均细胞丰度为2.49×103个/L,主要优势物种为菱形海线藻、伏氏海线藻(Thalassionema frauenfeldii)及短刺角毛藻(Chaetoceros messanensis)等;夏季调查区5m层浮游植物细胞丰度的平面分布由近岸到外海迅速减少,高值区主要在广东东部近岸及海南东部近岸;冬季则在珠江口近岸和海盆区出现较高值。两次调查中,浮游植物细胞丰度在浅水站位(200m)远高于深水站位(200m);较冬季相比,夏季浮游植物平均细胞丰度偏高,但物种丰富度却略偏低。夏、冬季浮游植物的香农-威纳多样性指数平均值分别为2.12和2.36,Pielou均匀度指数平均值分别为0.79和0.81。两个航次调查中,浮游植物细胞丰度均与盐度表现出显著性负相关性;在冬季还与磷酸盐浓度表现出显著性负相关性。     

南海北部和西菲律宾海颗石藻种群特征及差异

本文对南海北部SCS-N站(116°E,18°30′N,2016年5月)和菲律宾海西部PAC-ST02-34站(129°59.7′E,17°29.7′N,2016年1月)真光层中颗石藻种群结构进行了对比。总体而言,南海北部真光层颗石藻丰度高于西菲律宾海,而物种多样性指数低于西菲律宾海,这与南海北部真光层中营养盐浓度高于同纬度西菲律宾海有关;两个海域颗石藻丰度极大值都出现在次表层,但在南海北部出现的深度要浅于西菲律宾海,这是由于南海北部真光层营养盐、温、盐跃层较西菲律宾海更浅。Florosphera profunda是两个海域颗石藻种群的主要优势种,南海北部次级优势种为Emiliania huxleyi和Gephyrocapsa oceanica,而西菲律宾海次级优势种则为Umbellosphaera irregularis。颗石藻属种的垂直分布构成在两个海域具有相似的特征,其中F.profunda主要分布在真光层中深层(80m);G.oceanica和Gladiolithus flabellatus也主要出现在真光层中深层;Umbellosphaera irregularis,Syracosphaera pulchra,Helicosphaera carteri和Discosphaera tubifera等则主要分布在100m以浅;E.huxleyi和Calcidiscus leptoporus在南海北部主要分布在100m以浅,但在西菲律宾海则出现在深层(150m附近)。     

Incorporation of zinc into the coccoliths of the microalga <Emphasis Type="Italic">Emiliania huxleyi</Emphasis>

The coccolithophore Emiliania huxleyi is covered with elaborated calcite plates, the so-called coccoliths, which are produced inside the cells. We investigated the incorporation of zinc into the coccoliths of E. huxleyi by applying different zinc and calcium amounts via the culture media and subsequently analyzing the zinc content in the cells and the Zn/Ca ratio of the coccoliths. To investigate the Zn/Ca ratio of coccoliths built in the manipulated media, the algae have first to be decalcified, i.e. coccolith free. We used a newly developed decalcification method to obtain ‘naked’ cells for cultivation. E. huxleyi proliferated and produced new coccoliths in all media with manipulated Zn/Ca ratios. The cells and the newly built coccoliths were investigated regarding their zinc content and their Zn/Ca ratio, respectively. High zinc amounts were taken up by the algae. The Zn/Ca ratio of the coccoliths was positively correlated to the Zn/Ca ratio of the applied media. The unique feature of the coccoliths was maintained also at high Zn/Ca ratios. We suggest the following pathway of the zinc ions into the coccoliths: first, the zinc ions are bound to the cell surface, followed by their transportation into the cytoplasm. Obviously, the zinc ions are removed afterwards into the coccolith vesicle, where the zinc is incorporated into the calcite coccoliths which are then extruded. The incorporation of toxic zinc ions into the coccoliths possibly due to a new function of the coccoliths as detoxification sites is discussed.     

COCCOLITH PRODUCTION AND DETACHMENT BY EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE)1

Laboratory experiments were performed with the prymnesiophyte Emiliania huxleyi (Lohm.) Hay and Mohler, strain 88E, to quantify calcification per cell, coccolith detachment, and effects of coccolith production on optical scattering of individual cells. ¹⁴C incorporation into attached and detached coccoliths was measured using a bulk filtration technique. ¹⁴C-labeled cells also were sorted using a flow cytometer and analyzed for carbon incorporation into attached coccoliths. The difference between the bulk and flow cytometer analyses provided a ¹⁴C-based estimate of the rate of production of detached coccoliths. Coccolith production and detachment were separated in time in batch cultures, with most detachment happening well after calcification had stopped. Accumulation of coccoliths was maximum at the end of logarithmic growth with 50–80 coccoliths per cell (three to five complete layers of coccoliths around the cells). Net accretion rates of coccoliths were on the order of 7 coccoliths· cell^?1·d^?1 while net detachment rates were as high as 15 coccoliths· cell^?1·d^?1 for stationary phase cells. Equal numbers of coccoliths were attached and detached early in logarithmic growth, and as cells aged, the numbers of detached coccoliths exceeded the attached ones by a factor of 6. Our results demonstrate pronounced charges of forward angle light scatter and 90° light scatter of cells as they grow logarithmically and enter stationary phase. Counts of loose coccoliths in batch cultures are consistent with the detachment of coccoliths in layers rather than individual coccoliths.     

Movement of oceanic fronts south of Australia during the last 10 ka: interpretation of calcareous nannoplankton in surface sediments from the Southern Ocean

Preservation of calcareous nannoplankton in surface sediment samples from the Southern Ocean south of Australia and adjacent to New Zealand record a single assemblage. The dominant species are Emiliania huxleyi, Gephyrocapsa muellerae, Calcidiscus leptoporus, Helicosphaera carteri and Coccolithus pelagicus. The assemblage varies little in abundance and diversity with minor correlation to present-day overlying surface water masses and oceanic fronts. Increase in abundance of H. carteri and C. pelagicus in the region of the Subtropical Front may reflect higher nutrients associated with this front. The assemblage, although altered by dissolution, represents a warmer climatic interval than present-day with the presence of preferentially dissolved, warm-water species preserved as far south as the Polar Front. The presence of warm-water species under sub-Antarctic waters at the Polar Front is interpreted as a relic population from the Holocene climatic optimum of 10–8 ka. The absence of coccoliths in sediments poleward of the Polar Front suggests an equatorward shift of this front following the climatic optimum, resulting in increased productivity of siliceous phytoplankton associated with the colder waters and increased dissolution of coccoliths. Movement of the Subtropical Front for the same interval is not recorded in the preserved coccoliths. The more heavily calcified form of E. huxleyi which dominates the living assemblage north of the Subtropical Front is subject to dissolution in this region and is poorly preserved in the sediment assemblage.     

Distribution of large Emiliania huxleyi in the Central and Northeast Atlantic as a tracer of surface ocean dynamics during the last 25,000 years

The coccolithophore species Emiliania huxleyi is characterized by a wide range of sizes, which can be easily distinguished in the light microscope. In this study we have quantified the abundance of large (coccoliths > 4 µm in maximum length) E. huxleyi specimens during the last 25 kyr in sedimentary records from eleven cores and drill sites in the NE Atlantic and W Mediterranean Sea, to prove its usefulness in the reconstruction of water mass dynamics and biostratigraphic potential.During the Last Glacial Maximum this large form, a cold-water indicator, was common in the NE Atlantic and Mediterranean, and its regional variation in abundance indicates a displacement of the climatic zones southwards in agreement with the development of ice sheets and sea ice in the Northern Hemisphere during this period. On the other hand, the gradient between northern and southern surface water masses in the Subtropical Gyre appears to have been more pronounced than at present, while the Portugal and Canary Currents were more intense. In the western Mediterranean basin temperatures were cooler than in the adjacent Atlantic, provoking a quasi-endemism of these specimens until the end of Heinrich Event 1. This may have been due to a restriction in the communication between the Atlantic and Mediterranean through the Strait of Gibraltar, the arrival of cold surface water and the amplification of cooling after the development of ice sheets in the Northern Hemisphere.During the deglaciation, large E. huxleyi specimens decreased in abundance at medium and low latitudes, but were still numerous close to the Subarctic region during the Holocene. In transitional waters this decrease to present day abundances occurred after Termination Ib.The abrupt change in abundance of this large E. huxleyi form is proposed as a new biostratigraphic event to characterize the Holocene in mid- to low-latitude water masses in the North Atlantic, although this horizon seems to be diachronous by 5 kyr from tropical to subarctic regions, in agreement with the gradual onset of warm conditions.     

Mg isotopes and Mg/Ca values of coccoliths from cultured specimens of the species Emiliania huxleyi and Gephyrocapsa oceanica

Coccoliths from cultured specimens of two species of coccolithophores (Emiliania huxleyi and Gephyrocapsa oceanica) were sampled during two growth phases (late exponential and stationary), and their Mg isotope values (δ²⁶Mg) as well as Mg/Ca values were measured in order to investigate whether δ²⁶Mg can be used as a temperature proxy. Mg/Ca values were positively related with temperature (~ 0.002 mmol/mol/°C), without statistically significant differences between the two growth phases and the two species. Both species were depleted in heavier Mg isotopes relative to the culture medium, and δ²⁶Mg values were temperature dependent in both growth phases of E. huxleyi, although the δ²⁶Mg values differed in the two growth phases. In G. oceanica, a weak correlation between δ²⁶Mg values and temperature was seen in the late exponential growth phase only, and the δ²⁶Mg values differed between growth phases. The large differences between δ²⁶Mg values as measured in calcite formed during different growth phases indicate that Mg isotopes of coccoliths cannot be simply used as a temperature proxy. Our conclusions are preliminary and more data must be collected in order to fully evaluate the use of Mg isotopes of coccoliths as a temperature proxy.     

Patterns of carbon and nitrogen uptake during blooms of Emiliania huxleyi in two Norwegian fjords

Blooms of the coccolithophorid Emiliania huxleyi were monitoredin two land-locked fjords, Fauskangerpollen and Nordåsvannet(Western Norway), in May 1993. The chemical composition of particulatematter, size-fractionated photosynthesis, calcification, nitrogenuptake rates and the patterns of macromolecular synthesis wereexamined during the peak and decline of E.huxleyi blooms. Thetemporal evolution of the bloom in Fauskangerpollen was definedby a gradual decrease in cell abundance and cell-specific calcificationrates, and by increasing numbers of empty coccospheres and theratio detached coccoliths/living cells. A large proportion ofthe nitrogen required for microplankton growth was suppliedby aminonium and dissolved organic compounds such as urea and,as a consequence, the f-ratios were low (0.2). In general, nitrogenuptake patterns were consistent with ambient concentrationsof nitrogenous species. The photosynthetic carbon metabolismof E.huxleyi dominated phytoplankton assemblages was characterizedby high carbon allocation into lipids and relatively low carbonincorporation into protein as compared with diatom-dominatedassemblages. Consequently, the organic C/nitrogen uptake ratiodetermined stoichiometrically was significantly higher (up to10.8) when coccolithophorids were dominant than in diatom-basedor mixed-phyto-plankton assemblages. These carbon incorporationpatterns were reflected in differences in the chemical compositionof particulate matter.     

Induction of Phase Variation Events in the Life Cycle of the Marine Coccolithophorid Emiliania huxleyi

Emiliania huxleyi is a unicellular marine alga that is considered to be the world's major producer of calcite. The life cycle of this alga is complex and is distinguished by its ability to synthesize exquisitely sculptured calcium carbonate cell coverings known as coccoliths. These structures have been targeted by materials scientists for applications relating to the chemistry of biomedical materials, robust membranes for high-temperature separation technology, lightweight ceramics, and semiconductor design. To date, however, the molecular and biochemical events controlling coccolith production have not been determined. In addition, little is known about the life cycle of E. huxleyi and the environmental and physiological signals triggering phase switching between the diploid and haploid life cycle stages. We have developed laboratory methods for inducing phase variation between the haploid (S-cell) and diploid (C-cell) life cycle stages of E. huxleyi. Plating E. huxleyi C cells on solid media was shown to induce phase switching from the C-cell to the S-cell life cycle stage, the latter of which has been maintained for over 2 years under these conditions. Pure cultures of S cells were obtained for the first time. Laboratory conditions for inducing phase switching from the haploid stage to the diploid stage were also established. Regeneration of the C-cell stage from pure cultures of S cells followed a predictable pattern involving formation of large aggregations of S cells and the subsequent production of cultures consisting predominantly of diploid C cells. These results demonstrate the ability to manipulate the life cycle of E. huxleyi under controlled laboratory conditions, providing us with powerful tools for the development of genetic techniques for analysis of coccolithogenesis and for investigating the complex life cycle of this important marine alga.     

Seasonal variation in Emiliania huxleyi coccolith morphology and calcification in the Aegean Sea (Eastern Mediterranean)

A seasonal morphological variability is observed in Emiliania huxleyi var. huxleyi specimens, collected from discrete water samples in the Aegean Sea. Biometric analyses reveal a consistent pattern of increase in the size of coccoliths and coccospheres, including the thickness of the inner tube elements (INT), in winter/spring time low sea surface temperature and moderate productivity samples when compared with summer time high temperature-low productivity samples. The small range of salinity change in the Aegean Sea and the absence of seasonal pattern in nutrient content do not support any association with the observed increase in E. huxleyi coccolith size. A relatively increased [HCO₃^-] content is observed during spring-time interval related with the increase in the coccolith size, however it remains unclear which parameter of the carbonate system causes the observed effects.     

NEW EVIDENCE FOR MORPHOLOGICAL AND GENETIC VARIATION IN THE COSMOPOLITAN COCCOLITHOPHORE EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) FROM THE COX1b‐ATP4 GENES1

Emiliania huxleyi (Lohmann) W. W. Hay et H. Mohler is a cosmopolitan coccolithophore occurring from tropical to subpolar waters and exhibiting variations in morphology of coccoliths possibly related to environmental conditions. We examined morphological characters of coccoliths and partial mitochondrial sequences of the cytochrome oxidase 1b (cox1b) through adenosine triphosphate synthase 4 (atp4) genes of 39 clonal E. huxleyi strains from the Atlantic and Pacific Oceans, Mediterranean Sea, and their adjacent seas. Based on the morphological study of culture strains by SEM, Type O, a new morphotype characterized by coccoliths with an open central area, was separated from existing morphotypes A, B, B/C, C, R, and var. corona, characterized by coccoliths with central area elements. Molecular phylogenetic studies revealed that E. huxleyi consists of at least two mitochondrial sequence groups with different temperature preferences/tolerances: a cool‐water group occurring in subarctic North Atlantic and Pacific and a warm‐water group occurring in the subtropical Atlantic and Pacific and in the Mediterranean Sea.     

The mutual interplay between calcification and coccolithovirus infection

Two prominent characteristics of marine coccolithophores are their secretion of coccoliths and their susceptibility to infection by coccolithoviruses (EhVs), both of which display variation among cells in culture and in natural populations. We examined the impact of calcification on infection by challenging a variety of Emiliania huxleyi strains at different calcification states with EhVs of different virulence. Reduced cellular calcification was associated with increased infection and EhV production, even though calcified cells and associated coccoliths had significantly higher adsorption coefficients than non-calcified (naked) cells. Sialic acid glycosphingolipids, molecules thought to mediate EhV infection, were generally more abundant in calcified cells and enriched in purified, sorted coccoliths, suggesting a biochemical link between calcification and adsorption rates. In turn, viable EhVs impacted cellular calcification absent of lysis by inducing dramatic shifts in optical side scatter signals and a massive release of detached coccoliths in a subpopulation of cells, which could be triggered by resuspension of healthy, calcified host cells in an EhV-free, ‘induced media’. Our findings show that calcification is a key component of the E. huxleyi-EhV arms race and an aspect that is critical both to the modelling of these host–virus interactions in the ocean and interpreting their impact on the global carbon cycle.