免疫多糖对栉孔扇贝酸性磷酸酶、碱性磷酸酶和超氧化物歧化酶活性的影响

在栉孔扇贝的血清、血细胞和肝脏提取液中均发现有酸性磷酸酶（ＡＣＰ）、碱性磷酸酶（ＡＬＰ）和超氧化物歧化酶（ＳＯＤ）活性,其中,ＡＣＰ和ＡＬＰ在肝脏中的活性最高。采用１．０％的虫草多糖或海藻多糖作为免疫药物,对栉孔扇贝进行注射刺激后,发现血清中的ＡＣＰ、ＡＬＰ和ＳＯＤ活性有显著提高,血细胞中的ＡＣＰ和ＳＯＤ活性也有所增加,而肝脏提取液的这三种酶活性的变化不很明显。研究认为这两种多糖具有增强栉孔扇贝免疫活性的作用,在水产养殖业中有一定的开发价值。     

Alkaline phosphatase activity and its relationship to inorganic phosphorus in the transition zone of the North-western African upwelling system

The enzymatic activity of alkaline phosphatase (APA) was studied in the transition zone between the African upwelling system and the open ocean waters of the Canary Islands region. This region is recurrently dominated by the presence of upwelling filaments that may transport nutrient-enriched waters out into the open ocean before nutrients become exhausted by plankton. Turnover rates by APA were generally low in the whole region, but detectable in all the measurements carried out. On average, turnover rates were higher in the upwelling stations, and APA in those waters seemed to be mainly generated by heterotrophic bacteria to supply easily assimilable organic C. APA outside the upwelling area showed an inverse hyperbolic relationship with increasing phosphate, suggesting the presence of both constitutive and Pi-inducible APA. In these offshore waters, a threshold of 0.1 μM of phosphate could be defined for the regulatory function of Pi on APA. Thus, APA in nutrient-poor waters seemed to be induced to compensate for Pi-deficiency. Turnover rates in the filaments showed basal (probably constitutive) levels, whereas they increased in the surrounding waters, where phosphate concentration presumably did not satisfy plankton P-demands. The fertilising effect of the filaments and associated cyclonic eddies extended to at least 175 km offshore, where basal alkaline phosphatase activities were still found. The magnitude of this effect depends probably on the intensity of upwelling events and the degree of recirculation of filament water back to the coastal jet.     

海洋中碱性磷酸酶及其活性的研究进展

为加深对海洋生物地球化学循环过程和海洋生态系统功能的认知,并为相关研究提供科学依据,文章综述海洋中碱性磷酸酶及其活性的研究进展,并提出展望。研究结果表明：碱性磷酸酶主要来源于浮游植物、浮游动物和细菌,通常根据滤膜孔径等进行分类,对于海洋碳、氮、磷循环和调节生物群落组成具有重要意义;碱性磷酸酶活性通常采用水解荧光模拟底物法定量研究,并在全球海洋范围具有一定的水平、垂直和季节分布特征,影响碱性磷酸酶活性的典型环境因子包括太阳辐射、海水温度、河流输入、上升流、溶解态有机磷、溶解态无机磷、溶解态无机氮、金属离子以及病毒裂解、浮游动物捕食和排遗作用等。     

溶解态磷在海洋微藻碱性磷酸酶活力变化中的调控作用

在实验室批量培养条件下,测定了海洋微藻培养体系中碱性磷酸酶活力(APA)和各形态溶解磷的动态变化,分析了二者之间的关系.结果表明,在批量培养过程中,APA的动态变化呈"S"形曲线,各形态溶解磷在其变化过程中所起的调控作用不同,介质中溶解无机磷和小分子溶解有机磷的浓度是激发APA发生变化的主要调控因子,大分子溶解有机磷的浓度对APA的作用不明显,但APA的增大可加速微藻利用大分子溶解有机磷的速率.微藻的种类和丰度不影响APA的动态变化形式及其调控机理.     

中华绒螯蟹精子形成过程酸性磷酸酶的分布特征

将金属盐法与光镜、电镜技术相结合,研究中华绒螯蟹精子形成过程中酸性磷酸酶分布的变化。研究表明,在精子形成的早期,酸性磷酸酶由细胞质中的内质网小泡上产生,随后出现在细胞核内、顶体囊膜上及顶体管内,并且反应产物逐渐由分布均匀的细密颗粒聚集成分布均匀的较大的颗粒;当精子成熟后酸性磷酸酶均匀分布在顶体管中,反应产物颗粒比较大,穿孔器部分发现有少量酸性磷酸酶分布。     

Modern Dinoflagellate Cysts Distribution off the Eastern Part of Geoje Island,Korea

Distributional characteristics of dinoflagellate cysts in surface sediments were investigated in relation to environmental factors in the eastern part of Geoje Island, Korea. Samples were collected from 10 stations in February of 2004 and water temperature and salinity were measured in February, May, September and November of 2004. Total 30 taxa of dinoflagellate cysts were identified representing 19 genera, 28 species and 2 unidentified species. Among these dinoflagellate cysts,Brigantedinium spp. of which relative proportion in the total dinoflagellate cysts was 23.5%, was the most abundant at all stations except St. 1, and was followed bySpiniferites bulloideus (8.6%),Lingulodinium machaerophorum (8.2%) andDiplopsalis lenticula (6.7%). In addition, ellipsoidal cysts of the genusAlexandrium (Alexandrium catenella - tamarense type) andGymnodinium catenatum, known to be causative organisms for PSP, occurred with high concentrations.Scrippsiella trochoidea was also found; however, its cyst concentration was low. Generally, species composition in the study area was similar to these reported from Jinhae Bay and Busan Harbor and several dinoflagellate cysts reflected the eutrophic condition. Cyst distribution in the eastern part of Geoje Island seems to be influenced by the Tsushima Warm Current flowing from the southwest. The mean water temperature was 12.0°C in February, 14.7°C in May, 20.9°C in September and 17.2°C in November, which was most favorable forAlexandrium spp. growth. The abundances of dinoflagellate cysts ranged from 528 to 2,834 cysts/g dry sediment. Higher concentrations were recognized in sediments of west area of the Jisimdo than at other stations. The cyst composition of this area was closely related to these of Jinhae Bay and Busan Harbor from which currents flow into this area. Higher cyst concentration in the west area of Jisimdo might be due to formation of the gyre.     

大菱鲆仔稚鱼期消化酶及碱性磷酸酶活性的变化

针对海水鱼类仔稚鱼期高死亡率的现状,本实验采集不同发育阶段的大菱鲆,分析测定了酸、碱性蛋白酶、淀粉酶和脂肪酶以及消化道碱性磷酸酶的活性变化,以期为提高大菱鲆仔稚鱼期的成活率和生长率以及优化饵料配方提供理论依据。结果表明,大菱鲆初孵仔鱼体内就已具有碱性蛋白酶活性,而酸性蛋白酶活性在15日龄左右胃腺出现时才测得。蛋白酶活性的3个极低值分别出现于仔鱼开口期,卵黄囊完全消失期和仔鱼向稚鱼的转变期。早期仔鱼体内表现出较强的淀粉酶活性。5日龄时其活性急剧下降,并一直保持较低的比活力水平。脂肪酶活性在仔稚鱼期一直很弱,进入幼鱼期后比活力迅速升高。碱性磷酸酶活性从初孵到50日龄左右始终保持较小幅度的上升,直到60日龄变态完成时才出现大幅度的升高。大菱鲆仔稚鱼发育过程中,消化道中各种酶活性变化显著,仔稚鱼变态完成后,蛋白酶、脂肪酶和碱性磷酸酶活性显著增强,这标志着大菱鲆由稚鱼期向幼鱼期的转变及消化机能的完善。     

柘林湾近代沉积物中甲藻孢囊的垂直分布

于2001年4月在枯林湾3个采样点采集了各约20cm的沉积物柱状样,分层研究了甲藻孢囊的垂直分布,在33个沉积物榈中共分析鉴定出甲藻孢囊47种,而且还观察到了3种该海域特有的种类。孢囊组成以原多甲藻等异养型孢囊为主,而有毒的亚历山大藻和链状裸甲藻孢囊也均有分布。每个样品中所分析鉴定的孢囊种类数较少,仅有5-16种,平均10种,孢囊丰度也较低,在59-1998cysts/g D Wt之间,平均280cysts/g D Wt。香农-威弗种类多样性指数和孢囊种类数都以位于非养殖区湾外海域的S3站最高,同时在3个站位的6-8cm层次处,随深度的增加均有较大幅度的上升。研究结果表明,枯林湾的富营养化程度较高,而且湾内海域大于湾外海域,养殖区大于非养殖区,近年有加剧的趋势。     

夏季长江口不同粒级浮游植物碱性磷酸酶活性

碱性磷酸酶活性(alkaline phosphatase activity, APA)是海洋研究中用于反映浮游植物磷限制状态的重要指标。在长江口等“氮过剩”海域,磷是控制海域初级生产力水平的主要因子,然而,磷限制的范围常常难以界定,当前对不同粒级浮游植物的磷限制效应所知甚少。该文根据2020年夏季长江口航次资料,给出了海洋表层各粒级浮游植物(Net:≥20 μm;Nano:2~20 μm;Pico:0.8~2 μm)APA、浮游细菌APA(0.2~0.8 μm)和溶解态APA(<0.2 μm)的空间分布特征,并分析了APA与环境要素间的相关性。结果显示,各粒级浮游植物APA均与无机磷酸盐浓度(dissolved inorganic phosphate, DIP)呈显著负相关,这表明DIP浓度是影响浮游植物APA分布的主要因素。在平面分布上,各粒级浮游植物APA在近口门的光限制区均较低,且呈现自口门向外逐渐升高的趋势,与DIP的分布特征相反。Net和Nano级APA[平均值分别为 (40.28±32.35) nmol/(L·h)和(52.38±34.78) nmol/(L·h)]显著高于Pico级APA[平均值为(28.43±20.23) nmol/(L·h)],这表明大粒级浮游植物可能更易受DIP下降的影响。该研究中,诱导浮游植物APA快速升高的DIP浓度为0.159 μmol/L,与近岸区磷限制经验阈值相接近。该研究揭示了夏季不同粒级浮游植物对长江口磷分布的响应特征,有助于理解长江口初级生产过程的环境调控机制。     

Seasonal Blooms of the HAB Dinoflagellate Alexandrium taylori Balech in a New Mediterranean Area (Vulcano, Aeolian Islands)

Abstract. High-biomass blooms of A. taylori Balech have recently been spreading over new Mediterranean areas, with evident adverse effects on the marine ecosystem. In 1999–2000, a new Mediterranean locality was affected by blooms of A. taylori: the West Bay of Vulcano (Aeolian Islands, Tyrrhenian Sea), with maximum cell densities of 1.2 × 10⁷ cells 1⁻¹ in August 1999 and 4.0 × 10⁶ cells 1⁻¹ in August 2000 observed together with yellowish water discoloration. The seawater samples contained high concentrations of nutrients as DIN (Dissolved Inorganic Nitrogen), especially NH₃-N with values of 14.4 μM and TOT-P (Total Phosphorus) with values of 3.2 μM due to the anthropic presence and discharge of untreated sewage. The climatic conditions also seem to influence the occurrence and spreading of the A. taylori blooms in the Vulcano Bay. Clonal cultures of A. taylori , established from Italian and Spanish seawater samples, were used for the sequence analyses of the 5.8S rDNA gene and ITS regions in order to study the genetic variability of different geographical populations of Alexandrium species in the Mediterranean area and to further develop the molecular markers for HAB key-species.     

Alkaline phosphatase activity and phosphatase hydrolyzable phosphorus for phytoplankton in hiroshima bay, Japan

We investigated the seasonal variability of free alkaline phosphatase activity in seawater and alkaline phosphatase hydrolysable phosphorus (APHP) at 3 stations in Hiroshima Bay using alkaline phosphatase extracted from the dinoflagellates Alexandrium tamarense and Gymnodinium catenatum. The dissolved inorganic phosphorus (DIP) was lower than 1 μM in all samples; the lowest values were in May. The amount of APHP was high at the surface and bottom waters of all stations in May, showing DIP-depleted conditions. In August and November, the amount of APHP was much less than the amount of APHP in May, indicating that the availability of dissolved organic phosphorus (DOP) for these species was low and/or uptake during the dinoflagellate blooming might have occurred in the area. The results obtained from short-term variations of AP activity might suggest that the growth of dinoflagellates in this season may be partly supported by the AP produced by other diatoms.     

Use of Cell-Specific PAM-Fluorometry to Characterize Host Shading in the Epiphytic Dinoflagellate Gambierdiscus toxicus

Abstract. Cell‐specific fluorescence characteristics were used to characterize the light tolerance of the toxic benthic dinoflagellate Gambierdiscus toxicus. The fluorescence parameter F_v : F_m was measured using pulse amplitude modulation (PAM) fluorometry on individual cells collected from foliose red algae growing in the sub‐tidal margin of South Water Cay, Belize. Samples were collected over several days during sunny and cloudy conditions and compared to samples incubated in situ. The data from individual cells were used to generate both F_v : F_m frequency histograms and averages. Maximum individual cell values of F_v : F_m reached 0.81 in pre‐dawn samples, a value near the theoretical maximum for PAM fluorometry. In field samples from macroalgal hosts, average F_v : F_m values declined only slightly during the day, but cells incubated in bottles under 47 % incident sunlight showed a significant mid‐day depression. In freshly collected samples, near‐maximum F_v : F_m values could be found in individual cells during the entire day; however, the frequency histograms indicated a greater range in F_v : F_m values during the afternoon than in the morning. In contrast, cultures of G. toxicus showed a tight distribution around a mean. Field samples showed a rapid recovery to near‐maximum F_v : F_m within 2 min when assayed using a standardized actinic light series. Similar results were obtained in laboratory cultures of G. toxicus grown at 73 µmol photons · m^‐2 · s^‐1, but not at 383 µmol photons · m^‐2 · s^‐1. These data provide empirical support for suggestions that G. toxicus exploits the three‐dimensional structure of the algal host thallus to minimize light exposure. This strategy permits G. toxicus, a high‐light intolerant species in culture, to thrive in shallow, well‐lit tropical seas. It may also partially explain the observed preference of G. toxicus for complex, foliose macroalgae as hosts.     

毒死蜱和重金属对近海沉积物碱性磷酸酶和过氧化氢酶活性的影响

选择连云港近海临洪河口河心浅滩沉积物,采用不同浓度毒死蜱和重金属于室内培养,研究了毒死蜱和重金属对沉积物碱性磷酸酶和过氧化氢酶酶活性及其动力学特征的影响。研究结果表明:1)碱性磷酸酶活性受毒死蜱影响显著,峰值出现在10-200mg/kg浓度区间,培养第三周时V_max显著下降,酶活性抑制作用最显著。2)毒死蜱对过氧化氢酶激活作用的峰值出现在50mg/kg处。培养三周,过氧化氢酶活性逐渐回归正常值。3)Cd处理对沉积物碱性过氧化氢酶和过氧化氢酶抑制作用显著,明显强于Pb处理以及Cd、Pb复合处理,而Pb在相当程度上缓解了Cd对两种酶的抑制作用。4)酶活性可以用来指示沉积物的早期污染状况。     

Alkaline phosphatase activity of water column fractions and seagrass in a tropical carbonate estuary, Florida Bay

Few phosphorus-depleted coastal ecosystems have been examined for their ability to hydrolyze phosphomonoesters. We examined seasonal (August 2006–April 2007) alkaline phosphatase activity in Florida Bay, a phosphorus-limited shallow estuary, using fluorescent substrate at low concentrations (≤2.0 μM). In situ dissolved inorganic and organic phosphorus levels and phosphomonoester concentrations were also determined. Water column alkaline phosphatase activity was partitioned into two particulate size fractions (>1.2 and 0.2–1.2 μm) and freely dissolved enzymes (<0.2 μm). Water column alkaline phosphatase activity was also compared to leaf and epiphyte activity of the dominant tropical seagrass Thalassia testudinum. Our results indicate: (1) potential alkaline phosphatase activity in Florida Bay is high compared to other marine ecosystems, resulting in rapid phosphomonoester turnover times (2 h). (2) Water column alkaline phosphatase activity dominates, and is split equally between particulate and dissolved fractions. (3) Alkaline phosphatase activity was highest during cyanobacterial blooms, but not when normalized to chl a. These results suggest that dissolved, heterotrophic and autotrophic alkaline phosphatase activity is stimulated by phytoplankton blooms. (4) The dissolved alkaline phosphatase activity is relatively constant, while the particulate activity is seasonally and spatially dynamic, typically associated with phytoplankton blooms. (5) Phosphomonoester concentrations throughout the bay are low, even though potential hydrolysis rates are high. We propose that bioavailable dissolved organic P is hydrolyzed by dissolved and microbial alkaline phosphatase enzymes in Florida Bay. High alkaline phosphatase activity in the bay is also promoted by long hydraulic residence times. This background activity is primarily driven by carbon and phosphorus limitation of microorganisms, and regeneration of enzymes associated with cell lysis. Pulses of inorganic phosphorus and labile organic phosphorus and nitrogen may stimulate autotrophs, particularly cyanobacteria, which in turn promote biological activity that increase alkaline phosphatase activity of both autotrophs and heterotrophs in the bay.     

钦州湾磷营养状态与浮游植物的碱性磷酸酶活性分析

碱性磷酸酶(Alkaline phosphatase,AP)是浮游植物在磷胁迫的情况下诱导表达的一种酶,其作用是能够将溶解有机磷(Dissolved organic phosphate,DOP)水解成浮游植物可以直接利用的正磷酸盐形式,因此碱性磷酸酶活性(Alkaline phosphatase activity,APA)可用来指示海区浮游植物的磷胁迫状态。本研究以2015-2016年钦州湾4个航次(春季、夏季、秋季、冬季)的调查资料为依据,研究了钦州湾海域表层无机及有机磷的空间分布特征、APA的分布及其与磷营养之间的调控关系。调查期间,钦州湾海域春季、夏、秋、冬的单位叶绿素APA的平均值分别为20.42±5.32 nmol/μg Chl a/h,138.17±94.32 nmol/μg Chl a/h,142.60±72.60 nmol/μg Chl a/h和29.48±18.52 nmol/μg Chl a/h。经分析,APA与无机磷之间存在显著的负相关(P0.05),与N/P呈极显著负相关(P0.01),该海区APA可以反映钦州湾浮游植物的磷胁迫水平。海区氮磷比平均值在除春季外的三个季节均高于35,存在潜在的磷限制,浮游植物在夏季和秋季遭受较严重的磷胁迫。夏季和秋季浮游植物大量表达AP,由此推测DOP可能在钦州湾浮游植物生物量的维持甚至赤潮发生期间的磷补给上起着重要作用。     

Alkaline phosphatase in a littoral Mediterranean marine ecosystem: role of the main plankton size classes

This work investigates the alkaline phosphatase activity in a littoral marine ecosystem (Toulon Bay and Le Niel Bay, France) in order to study its biochemical characteristics with respect to pH, sea water composition and phosphate sensitivity. We also characterise the active forms in sea water and determine the extent to which zooplankton generate phosphatase activity with respect to other plankton classes. In Toulon Bay, phosphatase was produced mostly by the microplankton fraction (>90 μm), accounting for more than 90% of total activity. In contrast, most of the phosphatase activity in Le Niel Bay was generated by the nanoplankton fraction (5–90 μm) and the picoplankton fraction (0.25–5 μm). The microplankton enzymes had non Michaelis-Menten kinetics suggesting the involvement of multiple enzyme processes with distinct kinetic constants. This activity is in major part secreted into the sea water and is stimulated by the ionic strength and the pH of the sea water. Cypris larvae of the genus Balanus played a special role in this release. For the nanoplankton and picoplankton, part of this activity was due to non-secreted enzymes, probably bound to membranes or occurring intracellularly. Moreover, nano and picoplankton phosphatase required higher pH than microplankton enzyme. For all plankton size classes, there was no activity at low pH, suggesting that acid phosphatases were not involved in reactions with substrates dissolved in water.     

急性低盐度胁迫对可口革囊星虫存活、Na+/K+-ATP酶及磷酸酶活力的影响

为探讨低盐度急性胁迫对可口革囊星虫(Phascolosoma esculenta)的影响规律,研究可口革囊星虫对急性低盐度胁迫的响应机制,本试验先通过急性毒性试验确定可口革囊星虫96 h内对低盐度的耐受范围,再选取96 h最小死亡率盐度,研究该盐度急性胁迫下可口革囊星虫(平均体质量0.85±0.07 g)体质量、体壁含水量、Na~+/K~+-ATP酶活力以及酸碱磷酸酶活力在96 h内的动态变化。结果表明:在本试验盐度3.0～7.0范围内,可口革囊星虫24 h的最大死亡率盐度是3.0,可口革囊星虫96 h的最小死亡率盐度为7.0;可口革囊星虫死亡率随盐度降低逐渐升高,但相同盐度下随着时间延长,可口革囊星虫死亡率基本不变,可口革囊星虫96h内能耐受的最低盐度是7.0;在低盐度7.0胁迫下,可口革囊星虫体质量和体壁含水量在12 h内显著增大(P0.05),12 h后趋于平稳(P0.05);Na~+/K~+-ATP酶活力也在6 h内显著升高至最大值(P0.05),6 h至12h又显著降低(P0.05),此后稳定在高于对照组水平;酸性磷酸酶和碱性磷酸酶活力变化趋势与Na~+/K~+-ATP酶活力类似,均是先显著升高(P0.05)后显著降低(P0.05),最后稳定在高于对照组水平。以上研究表明,可口革囊星虫能够通过Na~+/K~+-ATP酶、酸碱磷酸酶等渗透和免疫相关的生理调节机制,迅速响应低盐度环境,其对低盐度有很强的适应力和耐受力。     

The use of surface alkaline phosphatase activity in the seagrass Posidonia oceanica as a biomarker of eutrophication

Eutrophication is one of the most relevant man‐induced changes occurring in coastal waters. The identification and assessment of specific responses to eutrophication in seagrasses can provide a useful tool for the detection of changes in the water quality in coastal zones, given the wide range of distribution of these organisms. In this study, we combine a correlational (across‐sites comparison) and a manipulative (fertilization experiment) approach to evaluate the usefulness and potential of alkaline phosphatase activity (APA) in the endemic Mediterranean seagrass Posidonia oceanica as an eutrophication biomarker. Our results showed that APA decreases promptly following nutrient additions, the response being maintained except during the winter period. APA also varies across natural meadows under different levels of nutrient discharges at scales relevant for monitoring purposes. AP activity seems to be an optimal ‘physiological biomarker’ that responds promptly and reliably to a pulse of eutrophication exposure. However, other considerations, such as the seasonality (the response disappears in winter), suggest its use with some caution and, as far as possible, as a complement of other bio‐indicators.     

中国对虾血淋巴抗菌活力的测定

采用溶藻胶弧菌（Ｖｉｂｒｉｏ ａｌｇｉｎｏｌｙｔｉｃｕｓ）、副溶血弧菌（Ｖ． ｐａｒａｈａｅｍ ｏｌｙｔｉｃｕｓ）、哈维氏弧菌（Ｖ．ｈａｒｖｅｙｉ）、嗜水气单胞菌（Ａｅｒｏｍ ｏｎａｓｈｙｄｒｏｐｈｉｌａ）对中国对虾（Ｐｅｎａｅｕｓ ｃｈｉｎｅｎｓｉｓ）血浆样品、血细胞溶解物（ＨＬＳ）样品的抗菌活力进行了测定。表明中国对虾血浆样品对四种细菌均表现出一定的抗菌活力。其中,三种弧菌活细胞数在孵育１ｈ 时呈现最低;嗜水气单胞菌活细胞数在孵育４ｈ 时最少。在三种弧菌中,血浆样品对哈维氏弧菌表现出的抗菌活力最强,而对副溶血弧菌表现出的抗菌活力最弱。血细胞溶解物样品对三种弧菌几乎没有表现出抗菌活力;嗜水气单胞菌在与血细胞溶解物样品孵育２ｈ 时,其活细胞数呈现最低,说明血细胞溶解物样品对该菌表现出了一定的抗菌活力。由此显示,抗菌活力在中国对虾血浆样品中表现得较强,而在血细胞溶解物样品中表现得较弱,血淋巴抗菌因子可以在短时间内对细菌表现出其最强的活力,但对不同种细菌其抗菌作用强弱不同。     

Effect of dissolved phosphorus on alkaline phosphatase activity in marine microalgae

I~IOXThe alkaline phOSPhataSe (AP) has been more and more studied following the study of thephosphorus limitation (Suttle and Harrision, 1988; Hu et al., 1989; Huang et al., 1996). PeOple used to take dissolved inorganic phOSphorus (DIP) level and uptake rate by algae as phosphOrus limitation indicators (Vera and Petterson 1992; Huang and HOng, 1994). But recent resultsindicate that diSSOlved organic phosphorus (DOP) and particulate organic phOSphorus (PP) couldalso be utilized b…