不同氮素形态配比对甘薯前期氮代谢的影响及其生理机制

以3种不同类型的甘薯(Ipomoea batatas(L.)Lam.)为实验材料,根据氮素的3种形态设置5个配比处理(N1～N5),分别在栽秧后15、25和35 d取样测定甘薯不同器官的氮含量、功能叶氮代谢酶活性变化以及酶调控基因表达情况.结果显示:在同一生育期,N4和N5处理铵态氮和硝态氮配施下植株氮素的积累量明显高...     

外源输入氮的有效性及形态对植物生长与生理影响的研究进展

氮(N)是控制植物结构和功能以及维持生态系统稳定的重要营养元素之一,外源输入氮素的有效性及形态的差异对植物的生长发育和生理特征产生显著的影响。由于全球气候变化和人类活动的干扰,大气氮沉降量日益增加,氮形态也发生改变,严重破坏植物的正常生长和生态系统的平衡稳定,已成为研究学者关注的热点问题。本文综述了不同氮输入水平和形态对植物的生长、光合作用、养分吸收以及代谢酶活性等方面的影响,归纳出：(1)适量氮输入能促进植物生长、光合作用和养分吸收能力,但当超出植物承受的阈值后,则对其产生抑制作用;(2)由于植物对氮素形态吸收偏好的差异,铵态氮(NH₄⁺-N)和硝态氮(NO₃^--N)对植物的生长、光合作用、养分吸收以及代谢酶活性的影响效果不同,且适宜铵硝配比相较于单一某种氮素添加对大多数植物的促进作用更显著。提出未来的研究方向应着重考虑4个方面：(1)开展大尺度的长期监测控制实验;(2)利用分子生物学技术深入探究氮形态对植物影响的微观机理;(3)重点关注土壤根际环境对植物根系氮素吸收的影响;(4)综合分析氮输入与...     

溶解氧对稻田土壤氮素转化及水稻氮代谢影响研究进展

溶解氧(dissolved oxygen,DO)在稻田氮素循环和水稻生长发育中发挥了重要作用,深入研究溶解氧对稻田氮素转化和水稻氮代谢的影响,对减少稻田氮素流失和提高水稻氮素利用具有重要意义。近年来,研究已阐明了溶解氧对土壤固氮作用、氨化作用、硝化作用和反硝化作用等生物化学过程以及土壤微生物多样性的影响;大量研究也表明,溶解氧在水稻氮素吸收与利用中起到了关键作用。本文系统介绍了溶解氧浓度对微生物群落结构和活性的影响及其与稻田不同形态氮素间转化的关系,综述了水稻根系生长特性、氮素吸收和同化及相关氨基酸代谢对溶解氧浓度响应机理的最新研究进展,并针对现阶段稻田氧营养研究提出了水稻生产中存在的问题及研究展望。     

氮素形态配比对菜用大豆籽粒膨大过程中氮碳同化的影响

以菜用大豆品种‘理想95-1'为试材,通过蛭石盆栽试验研究了氮素不同形态配比对菜用大豆籽粒膨大过程中碳氮代谢的影响.结果表明:营养液中增加适当的铵态氮比例(25%～50%)有利于菜用大豆的生长发育,菜用大豆植株和荚果干鲜重、干物率显著增加,尤以硝铵比为75∶25最为显著.在高比例的硝态氮(100%)或铵态氮(75%)处理下,菜用大豆籽粒的硝酸还原酶(NR)和谷氨酰胺合成酶(GS)活性分别显著升高,氮代谢显著增强,可溶性蛋白含量迅速增加,但同期的籽粒淀粉酶活性较低,可溶性糖和淀粉含量显著下降,碳代谢受到显著抑制.在硝铵比为75∶25时,菜用大豆籽粒氮代谢强度适中,同期碳代谢显著增强,籽粒可溶性糖和淀粉含量显著升高,能维持籽粒正常的生理代谢,有利于菜用大豆籽粒发育过程中营养物质的积累.可见,硝态氮和铵态氮配比能显著影响菜用大豆籽粒发育过程中的碳氮代谢,籽粒碳氮代谢与其产量密切相关,可通过调节硝铵比来获得理想菜用大豆产量.     

铁线莲属菝葜叶铁线莲组修订

对毛茛科Ranunculaceae铁线莲属Clematis的菝葜叶铁线莲组sect. Naraveliopsis进行了全面修订, 确定此组共含21种1亚种和1变种; 写出此组的分类学简史和地理分布, 并讨论了此组在铁线莲属中的系统位置; 将此组划分为3亚组, 写出分亚组、分种检索表, 以及各种植物的形态描述、地理分布、生长环境等, 并附有多数种的墨线图。根据对此组植物形态特征的分析, 观察到以下重要演化趋势: (1)叶从单叶演变到二回羽状复叶或二回三出复叶; (2)花从两性到单性, 从无退化雄蕊到有退化雄蕊; (3)雄蕊从无毛到有毛; (4)药隔突起从短(0.5-0.7 mm)到长(8.5-10 mm)。根据上述演化趋势,花两性、雄蕊被毛、退化雄蕊存在的荔波铁线莲亚组subsect. Liboenses(1种,特产贵州荔波)和花由两性变为单性的亚组subsect. Macgregorianae(2种,特产菲律宾)被认为是菝葜叶铁线莲组的进化群。在原始的菝葜叶铁线莲亚组subsect. Smilacifoliae(花两性,雄蕊无毛; 18种,广布亚洲热带地区)中,具单叶,花无退化雄蕊,药隔突起较短的菝葜叶铁线莲C. smilacifolia和滇南铁线莲C. fulvicoma被认为是较原始的种,而具三出复叶和退化雄蕊的C. vietnamensis和丝铁线莲C. loureiriana,以及具羽状复叶和长药隔突起(长达10 mm)的C. papillosa等3种则被认为是此亚组的进化种。自中南半岛北部山地向西经云贵高原南部至喜马拉雅东部山区集中分布有菝葜叶铁线莲组的13种植物,这一山区地带被认为是此组的分布中心。在此山区地带中,菝葜叶铁线莲的分布区和滇南铁线莲的分布区重叠部分的山区可能是此组的起源中心。     

短尾铁线莲叶绿素荧光特性研究

运用便携式光合气体分析系统(LI-6400)对短尾铁线莲(Clematis brevicaudata)的叶绿素荧光动力学曲线、淬灭分析、荧光光曲线以及荧光ACI曲线等生理特性进行测定和对比分析。结果表明:随光照时间的增加,短尾铁线莲的荧光参数ETR、qP和NPQ逐渐上升并在28 min左右达到稳定。最大荧光产量Fm′在叶片转入黑暗后逐渐上升,在52 min左右趋于稳定;NPQ则开始下降,在50 min左右达到稳定。PhiPS2和PhiCO2呈线性正相关(P<0.01)并随着光照强度的增加而减小。短尾铁线莲对CO2浓度的变化十分敏感,在50~200μmol.mol-1范围内,CO2浓度的增加可显著提高其PhiPS2和PhiCO2的值,增加CO2浓度是提高短尾铁线莲光合速率的有效途径。     

铁线莲属大叶铁线莲组修订

对毛茛科Ranunculaceae铁线莲属Clematis的大叶铁线莲组sect. Tubulosae进行了修订, 确定此组含9种2变种和3变型, 对此组的分类简史和地理分布做了介绍, 研究了此属大多数种的花粉形态, 写出此组组下分类群的形态特征、地理分布等, 并附有全部种的插图。此组9种被划分为2亚组, 其中原始群羽叶铁线莲亚组subsect. Pinnatae(有2种, 1种分布于中国河北和东北, 另1种产日本)在木质藤本习性、花构造、花粉形态(具3沟)等方面与威灵仙组sect. Clematis颇为近似, 区别主要在于萼片在开放初期近直立, 以后平展, 雄蕊花丝被毛, 此亚组可能源于威灵仙组。进化群大叶铁线莲亚组subsect. Tubulosae(有7种, 分布于我国东部、北部和台湾, 朝鲜和日本)为直立多年生草本、小亚灌木或小灌木, 花通常杂性, 萼片直立(花萼呈筒状, 稀呈坛状), 顶端或上部反曲, 雄蕊常有毛, 花粉通常具散孔, 只在1种(原始种大叶铁线莲C. heracleifolia)具3沟, 此亚组可能由羽叶铁线莲亚组演化而来。     

铁线莲属翅果铁线莲组修订

对毛茛科铁线莲属Clematis中单型的翅果铁线莲组sect .Pterocarpa进行了分类学修订,写出了此组及其惟一种,翅果铁线莲C.brachyura的分类学简史和形态描述,给出此种的插图。根据其体态及花构造近似亚洲东部的辣蓼铁线莲C.mandshuricaRupr .和圆锥铁线莲C.ternifloraDC.(二种均为威灵仙组sect .Clematis的成员) ,推测翅果铁线莲可能源自威灵仙组。     

畜禽粪便堆肥过程中氮素的损失与控制

堆肥是实现畜禽粪便处理及资源化利用的有效途径,然而畜禽粪便堆肥过程中氮素损失较严重,对氮素损失与控制的研究可为有效控制氮素损失提供理论基础和实践参考.本文简述了畜禽粪便堆肥过程中氮素转化及主要损失途径,总结了影响堆肥氮素损失的主要因素(包括堆肥物料初始特性、堆肥环境参数和堆肥工艺条件),并综述了氮素损失控制措施(调节C、N代谢,改变氮素形态,添加NH₃吸附剂和调节通风与控温措施)的研究进展,此外,对未来的研究方向进行了展望.     

氮素营养对苗期菘蓝叶中硝酸还原酶活性与矿质元素吸收的影响

采用正交试验设计,研究铵态氮、硝态氮和酰胺态氮3种氮素形态及其不同浓度配比对苗期菘蓝的单株干重、叶内的硝酸还原酶活性及矿质元素吸收的影响。结果显示:(1)影响苗期菘蓝单株干重的氮素形态依次为酰胺态氮>铵态氮>硝态氮。(2)不同氮素形态对叶片硝酸还原酶活性影响有差异,铵态氮影响最大,其次是硝态氮和酰胺态氮。(3)不同形态氮素配合施用后均能促进P、K、Ca、Mg、Cd、Mn、Cr、Sr 8种元素的吸收,但不利于Ni和Fe的吸收;元素吸收受铵态氮影响最大的矿质元素有K、Ba、Se、Ni、B、Si、Fe 7种元素,受硝态氮影响最大的元素有P、Cd、Ti、Al、Cu 5种元素,受酰胺态氮影响最大的元素有Na、Ca、Mg、Zn、Mo、Mn、Cr、Sr 8种元素。研究表明,不同形态氮素对苗期菘蓝吸收矿质元素的影响存在很大的差异,应注重酰胺态氮与无机的铵态氮、硝态氮的配合施用;适宜氮素形态及其配比能提高叶中硝酸还原酶的活性并促进矿质元素的吸收,从而有效地促进菘蓝的生长。     

不同氮素水平对油麦菜栽培效果的综合评价分析

为探讨不同氮素水平的硝态氮与铵态氮对油麦菜(Lactuca sativa L.)生长及品质的影响,自主设计并制作了适合蔬菜水培的立体管道栽培系统,配制了NO3-：NH4+=10：0,NO3-：NH4+=7：1,NO3-：NH4+=7：3的营养液,通过模糊数学原理,采用层次分析法(AHP)确定品质的权重值,综合评价了不同氮素水平对油麦菜的栽培效果。结果表明,以NO3-：NH4+=7：3营养液的栽培效果最好,其次是NO3-：NH4+=7：1。当营养液的NO3-：NH4+=7：3时,油麦菜的产量最高,叶片叶绿素、可溶性糖、可溶性蛋白质含量最高,硝酸盐含量最少,其综合品质最好。层次分析法对油麦菜不同氮素水平营养液的栽培效果的综合评价有较好的适用性,有助于获得南方设施管道栽培油麦菜最佳的营养液配方。     

北亚热带3种典型森林群落对降水中氮、磷、硫的截留及分配特征

森林群落在净化空气、截留沉降污染物、改善地表水质等方面具有重要作用。本研究以北亚热带地区3种典型森林群落（毛竹林、杉木林、青冈阔叶林）为研究对象,通过分析沉降污染物（NH₄⁺-N、NO₃^--N、NO₂^--N、TP和SO₄^2-）在大气降水、林内穿透雨、树干茎流、枯透水和地表径流中的浓度和通量变化特征,探讨不同森林群落对氮、磷、硫的截留净化作用和分配特征。结果表明,该区域大气降水中NH₄⁺-N、NO₃^--N、NO₂^--N、TP和SO₄^2-年均浓度分别为1.06、0.61、0.04、0.07、1.84 mg/L,其年均pH为5.88;各森林群落林冠层能够调升降雨的pH且全年稳定,对TP和NH₄⁺-N均有吸附作用,截留率分别为79.09%-84.68%和30.88%-69.36%;而枯落物层则是林下氮、磷、硫的主要释放源,对NH₄⁺-N、NO₃^--N、TP和SO₄^2-均具有淋溶作用;此外,由地表径流（输出）与大气降水（输入）的对比分析可知,各林地对沉降污染物中氮、磷、硫的截留率均超过98%;3种森林群落对沉降污染物中氮、磷、硫的截留能力依次为：青冈阔叶林 > 毛竹林 > 杉木林,阔叶林对沉降污染物的净化能力要高于毛竹林及针叶的杉木林。     

丛枝菌根真菌和施加不同形态氮肥对杉木幼苗养分吸收的影响

为了解丛枝菌根真菌(AMF)和不同形态氮对杉木(Cunninghamia lanceolata)生长和养分吸收的影响,以1 a生杉木幼苗接种摩西球囊霉(Glomus mosseae)和添加不同形态氮(NH₄⁺-N和NO₃^–-N),对其养分元素和生长状况的变化进行研究。结果表明,AMF显著提高了杉木的苗高和生物量,促进了杉木对N、P、K、Ca、Mg、Fe和Na的吸收,AMF对微量元素Fe、Na的促进作用总体上要强于大量元素K、Ca。与NO₃^–-N相比,AMF显著提高了NH₄⁺-N处理杉木的生物量、总C和N、Ca、Mg、Mn含量,而且这种显著性在叶中普遍高于根和茎。接种AMF可以促进杉木幼苗的生长和对养分元素的吸收,且添加NH₄⁺-N处理的促进作用要强于NO₃^–-N。     

马尾松组培苗对氮素形态的生长响应

马尾松属高氮需求树种,然而在苗木培育中马尾松对氮素,尤其是不同形态氮素的需求尚不明确.该文以马尾松组培苗为试验材料,采用基质培养方法,针对硝态氮、铵态氮两种氮素形态均分别设置了2、4、8、16 mmol·L-14个处理,以不添加氮素为对照,对苗木的高径生长、根构型参数(总根长、总表面积、总体积、平均直径和根尖数)以及生...     

不同氮处理下速生柳对水体氮的吸收、分配及生理响应

采用水培法,研究了旱柳苗在外源添加不同氮水平(贫氮、中氮、富氮、过氮)的铵态氮(NH+4-N)和硝态氮(NO-3-N)的生长、氮吸收、分配和生理响应。结果表明:一定范围氮浓度的增加能够促进旱柳苗的生长,但过量氮会抑制其生长,且NH+4-N的抑制作用大于NO-3-N;两种氮处理下,旱柳表现出对NH+4-N的吸收偏好,在同一氮水平时,旱柳各部位氮原子百分含量Atom%15N(AT%)、15N吸收量和来自氮源的N%(Ndff%)均为NH+4-N处理大于NO-3-N处理,且随着氮浓度的增加,差异增大,且在旱柳各部位的分布为根﹥茎﹥叶;2种氮素过量和不足均会对旱柳根和叶生理指标产生不同的影响,其中在过氮水平时,NH+4-N和NO-3-N处理下根系活力比对照减少了50.61%和增加了19.53%;在过氮水平时,NH+4-N处理柳树苗根总长、根表面积、根平均直径、根体积和侧根数分别对照下降了30.92%、29.48%、19.44%、27.01%和36.41%,NO-3-N处理柳树苗相应的根系形态指标分别对对照下降了1.66%、5.65%、1.49%、5.06%和25.72%。可见,高浓度NH+4-N对旱柳苗的胁迫影响大于NO-3-N,在应用于水体氮污染修复时可通过改变水体无机氮的比例,削弱其对旱柳的影响,从而提高旱柳对水体氮污染的修复效果。     

根际不同供氮水平对紫青菜生长和营养品质的影响

为了解根际供氮对紫青菜(Brassica campestris ssp.chinensis var.communis)生长和营养品质的影响,用不同水平的NH_4NO_3处理,对紫青菜的生理特性和营养品质进行研究。结果表明,随着供氮水平的增加,紫青菜的鲜质量、干质量、根系活力和游离氨基酸含量均呈先上升后略下降趋势;硝酸还原酶(NR)活性、叶绿素含量、NO_3~–含量和可溶性蛋白质含量均呈上升趋势;DPPH·自由基清除率、超氧化物歧化酶(SOD)活性、过氧化物酶(POD)同工酶和过氧化氢酶(CAT)同工酶活性等均提高;而花青素苷相对含量、可溶性糖含量、维生素C(Vc)含量、总酚含量和FRAP值均呈下降趋势。根际NH_4NO_3为75~300 mg L~(–1)时,紫青菜的鲜质量和干质量较低;NH_4NO_3达1200 mg L~(–1)时,其鲜质量和干质量又低于600 mg L~(–1)处理的,植株积累较多NO_3~–,导致营养品质下降,同时根际也积累NH4NO3。因此,600 mg L~(–1)的根际NH_4NO_3是适宜的氮水平,其鲜质量和干质量均最高、营养均衡、抗氧化能力强。     

Occurrence and intensity of wild boar disturbances, effects on the physical and chemical soil properties of alpine grasslands

Background and aims

Physical and chemical soil properties determine local plant conditions and resources, affecting plants’ ability to respond to disturbances. In alpine grasslands, wild boar disturbances occur at different intensities, what may affect differently their soil properties. Alpine soils from five contrasted plant communities were explored within and outside disturbances, accounting for an overall and community scale effect. Additionally, we analysed the effect of disturbance intensity on soil NO₃ ^--N and NH₄ ⁺-N.

Methods

Soils were analyzed for physical (bulk density, moisture content and electrical conductivity), and chemical properties (pH, total N and C, oxidizable C, C:N ratio, available K, P, Ca²⁺, Na⁺ and Mg²⁺). Resin bags were used to compare the effect of the disturbance occurrence and intensity on soil NO₃ ^--N and NH₄ ⁺-N.

Results

Bulk density, total N and NO₃ ^--N concentration were significantly higher in disturbed areas, while soil moisture, C:N, NH₄ ⁺-N, Na⁺, Mg²⁺ and Ca²⁺ concentrations were significantly lower. However, low disturbance intensity reduced NO₃ ^--N and increased NH₄ ⁺-N concentrations.

Conclusions

Wild boar occurrence and intensity strongly alter physical and chemical conditions of alpine soils, increasing soil compaction, and altering the availability of N forms. These changes may affect most plant species, thus affecting the structure and dynamics of alpine plant communities.     

Nitrogen isotope discrimination in white spruce fed with low concentrations of ammonium and nitrate

Differences in ¹⁵N among ten white spruce [Picea glauca (Moench) Voss] families were examined in hydroponic experiments testing (1) three N sources [100 M N as (i) NH₄⁺, (ii) NO₃^– or (iii) NH₄NO₃] and (2) two supply regimes [200 M NH₄⁺ (i) maintained steadily or (ii) recurrently drawn-down]. In the N-source experiment, the NH₄⁺ treatment resulted in superior growth and lower C/N ratios. Whole plant ¹⁵N was higher on NH₄⁺ and NH₄NO₃, reflecting higher NH₄⁺ removal rates from the media. Families expressed differences in biomass, C/N, ¹⁵N and ¹³C. Family ¹⁵N and ¹³C were positively correlated in the NH₄NO₃ treatment and the steady-state regime. Supply regime did not affect total biomass, but higher root/shoot ratios implied N was more limiting under the draw-down regime. Family rank changed with supply regime, but not with N source. Analysis of media isotope enrichment during substrate depletion revealed relationships between net discrimination and external N concentration. Discrimination against ¹⁵NH₄⁺ was about twice that of ¹⁵NO₃^–. A simple model relating isotope discrimination to relative rates of ion efflux and influx predicted efflux/influx ratios consistent with published values for white spruce. We propose that genetic differences in discrimination are caused by different demands on assimilation and in uptake capacity which interact, influencing the balance between N influx and efflux.     

Growth,chemical composition,and nitrate reductase activity of Rumex species in relation to form and level of N supply

Growth, chemical composition, and nitrate reductase activity (NRA) of hydroponically cultured Rumex crispus, R. palustris, R. acetosa, and R. maritimus were studied in relation to form (NH₄ ⁺, NO₃ ^-, or both) and level of N supply (4 mM N, and zero-N following a period of 4mM N). A distinct preference for either NH₄ ⁺ or NO₃ ^- could not be established. All species were characterized by a very efficient uptake and utilization of N, irrespective of N source, as evident from high concentrations of organic N in the tissues and concurrent excessive accumulations of free NO₃ ^- and free NH₄ ⁺. Especially the accumulation of free NH₄ ⁺ was unusually large. Generally, relative growth rate (RGR) was highest with a combination of NH₄ ⁺ and NO₃ ^-. Compared to mixed N supply, RGR of NO₃ ^-- and NH₄ ⁺-grown plants declined on average 3% and 9%, respectively. Lowest RGR with NH₄ ⁺ supply probably resulted from direct or indirect toxicity effects associated with high NH₄ ⁺ and/or low Ca²⁺ contents of tissues. NRA in NO₃ ^- and NH₄NO₃ plants was very similar with maxima in the leaves of ca 40 μmol NO₂ ^- g^-1 DW h^-1. ‘Basal’ NRA levels in shoot tissues of NH₄ ⁺ plants appeared relatively high with maxima in the leaves of ca 20 μmol NO₂ ^- g^-1 DW h^-1. Carboxylate to organic N ratios, (C-A)/N_org, on a whole plant basis varied from 0.2 in NH₄ ⁺ plants to 0.9 in NO₃ ^- plants. After withdrawal of N, all accumulated NO₃ ^- and NH₄ ⁺ was assimilated into organic N and the organic N redistributed on a large scale. NRA rapidly declined to similar low levels, irrespective of previous N source. Shoot/root ratios of -N plants were 50–80% lower than those from +N plants. In comparison with +N, RGR of -N plants did not decline to a large extent, decreasing by only 15% in -NH₄ ⁺ plants due to very high initial organic-N contents. N-deprived plants all exhibited an excess cation over anion uptake (net proton efflux), and whole-plant (C-A)/N_org ratios increased to values around unity. Possible difficulties in interpreting the (C-A)/N_org ratio and NRA of plants in their natural habitats are briefly discussed.     

Nitrogen utilization by the raphidophyte Heterosigma akashiwo: Growth and uptake kinetics in laboratory cultures

The nitrogen uptake and growth capabilities of the potentially harmful, raphidophycean flagellate Heterosigma akashiwo (Hada) Sournia were examined in unialgal batch cultures (strain CCMP 1912). Growth rates as a function of three nitrogen substrates (ammonium, nitrate and urea) were determined at saturating and sub-saturating photosynthetic photon flux densities (PPFDs). At saturating PPFD (110 μE m⁻² s⁻¹), the growth rate of H. akashiwo was slightly greater for cells grown on NH₄⁺ (0.89 d⁻¹) compared to cells grown on NO₃⁻ or urea, which had identical growth rates (0.82 d⁻¹). At sub-saturating PPFD (40 μE m⁻² s⁻¹), both urea- and NH₄⁺-grown cells grew faster than NO₃⁻-grown cells (0.61, 0.57 and 0.46 d⁻¹, respectively). The N uptake kinetic parameters were investigated using exponentially growing batch cultures of H. akashiwo and the ¹⁵N-tracer technique. Maximum specific uptake rates (V_max) for unialgal cultures grown at 15 °C and saturating PPFD (110 μE m⁻² s⁻¹) were 28.0, 18.0 and 2.89 × 10⁻³ h⁻¹ for NH₄⁺, NO₃⁻ and urea, respectively. The traditional measure of nutrient affinity—the half saturation constants (K_s) were similar for NH₄⁺ and NO₃⁻ (1.44 and 1.47 μg-at N L⁻¹), but substantially lower for urea (0.42 μg-at N L⁻¹). Whereas the α parameter (α = V_max/K_s), which is considered a more robust indicator for substrate affinity when substrate concentrations are low (<K_s), were 19.4, 12.2 and 6.88 × 10⁻³ h⁻¹/(μg-at N L⁻¹) for NH₄⁺, NO₃⁻ and urea, respectively. These laboratory results demonstrate that at both saturating and sub-saturating N concentrations, N uptake preference follows the order: NH₄⁺ > NO₃⁻ > urea, and suggests that natural blooms of H. akashiwo may be initiated or maintained by any of the three nitrogen substrates examined.