Quality of micropropagated plants—Vitrification

Summary Vitrification-Hyperhydrous shoot development, effects the survival and quality of several micropropagated plants ex-vitro. The leaves which are the immediate organ to be affected, exhibit abnormal morphology and physiology. Leaf malfunction is apparently a stress response to very rich media and high relative humidity. The understanding of the underlying mechanism of vitrification and its control in vitro can contribute to a more efficient micropropagation. Vitrification was found to be associated with elevated ethylene production which was related to hypolignification and poor cell wall development. Liquid and low agar media induced callose formation along with reduced and disoriented cellulose biosynthesis, manifested also in non-functioning guard cells. Malfunctioning stomata, in addition to defective cuticle contributed to increased transpiration and desiccation of in vitro formed leaves. The activity of various enzymes, associated with cell wall synthesis, was low and total proteins in normal leaves was higher than in vitreous ones. Various measures were found to reduce vitrification; lowered matrix and water potential in the medium, reduction in RH, low NH ₄ ⁺ , changes in Ca⁺⁺ levels and the removal of ethylene. These measures improved leaf morphogenesis, survival and the quality of several micropropagated plant species. Presented in the Session-in-Depth Transition of Plants From Culture to Establishment In Vivo,“ at the 41st Annual Meeting of the Tissue Culture Association, Houston, Texas, June 10–13, 1990.     

黄河三角洲河滩与潮滩芦苇对盐胁迫的生理生态响应

研究黄河三角洲两种生境芦苇对盐胁迫的生理生态响应差异,能为退化滨海湿地生态修复中芦苇植株来源的选择提供重要的理论支持。在盐胁迫(300 mmol/L NaCl)下,比较研究了黄河三角洲河滩芦苇(低盐生境)和潮滩芦苇(高盐生境)叶片中的Na⁺含量、根部分生区Na⁺流速、叶片的光合作用参数、H₂O₂的含量、抗氧化酶的活性、丙二醛和脯氨酸的含量。结果表明:盐胁迫显著提高了河滩芦苇叶片中Na⁺含量,但对潮滩芦苇叶片Na⁺影响不显著。进一步通过非损伤微测技术研究发现,盐胁迫后,潮滩芦苇比河滩芦苇的根部分生区的Na⁺外排流速更高(潮滩芦苇:(1982.05±122.74) pmol cm^-2 s^-1vs.(87.93±12.94) pmol cm^-2 s^-1,P<0.01;河滩芦苇:(1574.16±458.90) pmol cm^-2     

Light and dark uptake and loss of 14C: methodological problems with productivity measurements in oceanic waters

Measurements of the uptake and loss of ⁴C in the light and in the dark in the Tasman and Coral Seas have revealed methodological problems with the estimation of productivity in these waters. Rates of productivity estimated without replication, time series incubations and dark controls frequently overestimated the true rates of autotrophic production. The data showed unexpectedly high rates of both uptake and loss in the dark in oligotrophic waters. In oligotrophic oceanic waters, dark incorporation of ¹⁴C sometimes equalled the uptake of ¹⁴C in the light bottle. Rapid uptake of isotope in the dark controls appeared to be the result of rapid bacterial growth and metabolism. This problem was exacerbated by agitation of the sample before or during the incubation. Tropical samples were particularly susceptible to problems arising from the agitation of the samples. Latitudinal gradients of dark uptake and loss were revealed in these incubations. The loss of label during 8–12 hours in the dark (after 12 hr in the light) was as high as 50% in subtropical waters. The loss was frequently unmeasurable (< 10%) in temperate waters. The time course of ¹⁴C uptake indicated active grazing in the bottles and suggested that most of the nighttime losses of label were due to grazing by microheterotrophs. Respiratory losses appeared to be small. Calculated values of the assimilation number (or photosynthetic capacity) which did not correct for dark ¹⁴C uptake were too high to be biochemically realistic. The errors were due to the heterotrophic uptake of label and the lack of dark controls. Rapid release of ¹⁴C in the dark after incubation in the light meant that the estimate of productivity was dependant on the trophic state of the sample and on the period of incubation.     

The role of the external mycelial network of vesicular-arbuscular mycorrhizal fungi: a study of carbon transfer between plants interconnected by a common mycelium

The transfer of ¹⁴C from Lolium perenne (the donor) to Plantago lanceolata (the receiver), mediated by vesicular-arbuscular (VA) mycorrhizal fungi, was examined when the two species were grown together or separately. The VA mycorrhizal infection led to a significant increase, relative to that in uninfected plants, in the ¹⁴C transferred from donor to receiver plants, not only when the roots of the two plants were growing in intimate mixture, but also when they were separated by a root-free zone of 2.33 cm. The majority of isotope transfer between the two plant species was along the direct pathway via VA mycelium.     

Effect of vertical cycling on photosynthesis during the annual algal succession in the northern Baltic

During the phytoplankton succession in 1984 and 1985, the effect of fluctuating light on algal photosynthesis (incorporation of ¹⁴C, acidified water sample) was studied in the northern Baltic. Bottles were mounted on moving racks that mimicked vertical transport caused by Langmuir circulations in the trophogenic layer. Assuming that the photoinhibition observed near the surface in fixed-depth incubations (from 1 to 8% of integral photosynthesis) was avoided in cycled samples, vertical cycling conducted around noon resulted in on average 10% lower photosynthesis than fixed-depth incubations (n = 17). This difference lies within the 5% confidence limits of the measurement, and hence it was concluded that the lack of short-term fluctuations in light associated with the vertical circulation of natural phytoplankton communities does not seriously bias conventional in situ ¹⁴CO₂ fixation measurements performed at fixed depths in the study area.     

Physiological responses of forest trees to heat and drought

The heat wave of summer 2003 was the largest and the most persistent ever experienced in Central Europe and has fuelled concern about the effects of climate change on European ecosystems. Since forests constitute the most important European ecosystems, in this review article we assess current knowledge on the effects of heat and drought on key metabolic processes for growth and productivity of forest trees. In particular, the general consequences of heat and drought on (1) photosynthesis and respiration at the cellular and community level, and (2) on nutrient uptake, partitioning and competition for nutrients are summarized. The latter are a major sink for photosynthetic energy and, therefore, are indirectly but strongly connected to the performance of photosynthesis. In addition, the interaction of heat and drought with stress compensation mechanisms and emission of biogenic volatile organic compounds (BVOC) are discussed, since these processes are directly connected to carbon metabolism. Effects on the emission of BVOC are also included because they constitute an important feedback mechanism on ozone formation and, thus, on atmospheric pollution. As far as available, data collected during the 2003 heat wave are included and discussed.     

酸胁迫下琥珀酸放线杆菌的生理及转录应答

【目的】通过琥珀酸放线杆菌Actinobacillus succinogenes CGMCC1593对酸胁迫的生理应答和转录组学分析,探究琥珀酸放线杆菌酸胁迫的机制。【方法】测定不同pH对细胞生长、H+-ATPase、细胞内pH的影响;测定酸胁迫前后细胞膜和谷氨酸脱氢酶的变化、谷氨酸对琥珀酸放线杆菌生长的影响;通过RNA-seq测序分析酸胁迫条件下的差异表达基因。【结果】随pH值的降低,细胞生长受抑制,H+-ATPase的活性下降。pH 4.7酸胁迫后,细胞膜受到严重损伤,谷氨酸对酸胁迫后的细胞有保护作用,GDH酶活响应酸胁迫后略有增加。酸胁迫后,39个基因差异表达较为显著,其中49%基因属于应激蛋白、转运蛋白,小部分基因与代谢相关。【结论】本文探究了琥珀酸放线杆菌酸胁迫下的生理及转录应答,研究结果可为寻找增强琥珀酸放线杆菌耐酸性策略提供参考。     

Physiological responses of wheat seedlings to drought and UV-B radiation. Effect of exogenous sodium nitroprusside application

Physiological and biochemical responses of wheat seedlings to drought, UV-B radiation, and combined stress were investigated. Drought, UV-B, and combined stresses retarded seedling growth by 26.5, 29.1, and 55.9%, respectively. One reason for growth retardation may be the oxidative damage indicated by an increase in the H₂O₂ content and lipid peroxidation degree. Furthermore, there was negative correlation between shoot fresh weight and H₂O₂ content, fresh weight and the content of thiobarbituric acid-reacting substances (TBARS), and the positive correlation between H₂O₂ content and TBARS (R ² = 0.9251, 0.9005, and 0.9007, respectively). The activities of superoxide dismutase, guaiacol peroxidase, and ascorbate peroxidase increased under drought, UV-B, and the combination of stresses, while catalase activity decreased under the combined stress as compared to the control. The combination of drought and UV-B caused more severe damage to wheat seedlings than stress factors applied separately. Thus, the combined application of drought and UV-B had more strong adverse effects on wheat seedlings. The addition of 0.2 mM sodium nitroprusside (SNP) enhanced wheat seedling growth under drought, UV-B, and combined stress, likely, due to decreasing the accumulation of H₂O₂ and lipid peroxidation as well as activating the antioxidant enzymes. However, SNP treatment decreased the proline content. Published in Russian in Fiziologiya Rastenii, 2007, Vol. 54, No. 5, pp. 763–769. The text was submitted by the authors in English.     

Characterization and identification of bacteria isolated from micropropagated mint plants

Summary Bacterial isolates from contaminated mint shoot cultures were characterized and identified as a preliminary step in determining an elimination treatment. The 22 bacteria were characterized using biochemical and morphological tests and subjected to sensitivity tests with four antibiotics. The isolates were compared with known organisms and assigned to genera according to similarities in characteristics. Seven isolates were analyzed by fatty acid analysis carried out by a commercial laboratory. Six were classified asAgrobacterium radiobacter; eight asXanthomonas; one each asPseudomonas fluorescens, Micrococcus spp.,Corynebacterium spp., andCurtobacterium spp.; four could not be assigned to genera. Inhibition of growth of the bacteria by most antibiotics was best at pH 7.5. Minimal inhibitory concentration and minimal bactericidal concentrations of gentamicin, rifampicin, streptomycin sulfate, and Timentin varied with genotype.     

Effect of rice plants on methane production and rhizospheric metabolism in paddy soil

In order to elucidate the effects of rice plants on CH₄ production, we conducted experiments with soil slurries and planted rice microcosms. Methane production in anoxic paddy soil slurries was stimulated by the addition of rice straw, of unsterile or autoclaved rice roots, and of the culture fluid in which rice plants had axenically been cultivated. The addition of these compounds also increased the concentrations of acetate and H₂, precursors of CH₄ production, in the soil. Planted compared to unplanted paddy soil microcosms exhibited lower porewater CH₄ concentrations but higher CH₄ emission rates. They also exhibited higher sulfate concentrations but similar nitrate concentrations. Concentrations of acetate, lactate and H₂ were not much different between planted and unplanted microcosms. Pulse labeling of rice plants with¹⁴CO₂ resulted during the next 5 days in transient accumulation of radioactive lactate, propionate and acetate, and after the second day of incubation in the emission of¹⁴CH₄. Most of the radioactivity (40–70%) was incorporated into the above-ground biomass of rice plants. However, during a total incubation of 16 days about 3–6% of the applied radioactivity was emitted as¹⁴CH₄, demonstrating that plant-derived carbon was metabolized and significantly contributed to CH₄ production. The sequence of the appearance of radioactive products and their specific radioactivities indicate that CH₄ was produced from root exudates by a microbial community consisting of fermenting and methanogenic bacteria.     

Validation of the design of feeding experiments involving [(14)C]substrates used to monitor metabolic flux in higher plants

The aim of this study was to examine whether flux through the pathways of carbohydrate oxidation is accurately reflected in the pattern of ¹⁴CO₂ release from positionally labelled [¹⁴C]substrates in conventional radiolabel feeding studies. Heterotrophic cell suspension cultures of Arabidopsis thaliana were used for this work. The presence of an alkaline trap to capture metabolically generated ¹⁴CO₂ had no significant effect on the ratio of ¹⁴CO₂ release from specifically labelled [¹⁴C]substrates, or on the metabolism of [U-¹⁴C]glucose by the cells. Although the amount of ¹⁴CO₂ captured in a conventional time-course study was only about half of that released from a sample acidified at an equivalent time point, the ratios of ¹⁴CO₂ released from different positionally labelled [¹⁴C]glucose and [1-¹⁴C]gluconate were the same in untreated and acidified samples. Less than 5% of radioactivity supplied to the growth medium as [¹⁴C]bicarbonate was incorporated into acid-stable compounds, and there was no evidence for appreciable reassimilation of ¹⁴CO₂ generated intracellularly during oxidation of [1-¹⁴C]gluconate by the cells. It is concluded that the ratio of label captured from specifically labelled [¹⁴C]glucose is a valid and convenient measure of the relative rates of oxidation of the different positional carbon atoms within the supplied respiratory substrate. However, it is argued that failure to compensate for the incomplete absorption of ¹⁴CO₂ by an alkaline trap may distort estimates of respiration that rely on an absolute measure of the amount of ¹⁴CO₂ generated by metabolism.     

Effect of Drought Stress on Certain Morphological and Physiological Characteristics of a Resistant and a Sensitive Canola Cultivar

Water stress is one of the main abiotic factors that reduces plant growth, mainly due to high evaporative demand and low water availability. In order to evaluate the effects of drought stress on certain morphological and physiological characteristics of two canola cultivars, we conducted a factorial experiment based on a completely randomized design. The findings show that drought stress exacerbations result in the plant's response to stress due to increased canola resistance caused by changes in plant pigments, proline, catalase, ascorbate peroxidase, peroxidase, superoxide dismutase and malondialdehyde, glucose, galactose, rhamnose and xylose. These in turn ultimately influence the morphological characteristics of canola. Drought stress reduces the concentration of carotenoids, chlorophyll a, chlorophyll b, total chlorophylls; however, glucose, galactose, rhamnose, xylose, proline, catalase, ascorbate peroxidase, peroxidase, superoxide dismutase, malondialdehyde (in leaves and roots) and the chlorophyll a and b ratios were increased. Reduction of plant height, stem height, root length, fresh and dry weight of canola treated with 300 g/l PEG compared to non‐treatment were 0.264, 0.236, 0.394, 0.183 and 0.395, respectively. From the two canola cultivars, the morphological characteristics of the NIMA increased compared to the Ks7 cultivar. Interaction effects of cultivar and drought stress showed that NIMA cultivar without treatment had the highest number of morphological characteristics such as carotenoid concentration, chlorophyll a, chlorophyll b, total chlorophylls a and b, whereas the cultivar with 300 g/l PEG (drought stress) had the highest amount of proline, malondialdehyde, soluble sugars and enzymes in leaves and roots. Increasing activity of oxidative enzymes and soluble sugars in canola under drought stress could be a sign of their relative tolerance to drought stress.     

Regulation of the utilization of glucose and aromatic substrates in four strains of Pseudomonas putida

During the oxidation of various mixtures of glucose and aromatic substrates by four strains of Pseudomonas putida, diauxic growth was not observed. Strain A3.12 grew faster on benzoate than on glucose, whereas three other strains showed faster growth on glucose than on the aromatic test substrates. Growth rates on mixtures of glucose and aromatics were intermediate between those on the single substrates.The presence of glucose in media containing aromatic substrates accelerated in the bacteria the appearance of the ability to oxidize aromatic substrates. During growth of the organisms on binary mixtures of aromatics, simultaneous utilization of these compounds occurred, the utilization ratio depending on the quality of the compounds as carbon and energy sources. Addition of glucose to dual aromatic substrate media greatly increased the utilization ratio in favour of the better aromatic substrate.With decreasing concentration of glucose in relation to that of aromatic substrates, the rate of carbon assimilation from glucose increased. Enzymological and radiochemical studies demonstrated that even in the presence of an excess of aromatic substrates, glucose was exclusively catabolized via the 2-keto-3-deoxy-6-phosphogluconate pathway. In contrast, the rate of carbon assimilation from ¹⁴C-ring-labelled benzoate and anisate was unaffected by the presence of an excess of glucose.Abbreviations KDPG 2-keto-3-deoxy-6-phosphogluconate - PP pentose-phosphate - OD optical density     

不同品种万寿菊对镉胁迫的生长和生理响应

以6个万寿菊品种为研究对象,研究了不同浓度的重金属镉（Cd）胁迫（0、200、400、600 mg/L）对万寿菊基径、株高以及百叶重、茎叶重和单花质量等生长指标及其质膜透性（丙二醛、相对电导率）、保护酶活性（SOD、POD、CAT、APX）以及光合特性(叶绿素含量、净光合速率、气孔导度、蒸腾速率、水分利用效率)等生理指标的影响。结果显示:(1)低于400 mg/L的Cd胁迫对6个万寿菊品种的基径、株高无明显影响,而百叶重、茎叶重和单花质量却随着Cd浓度的增加显著降低,尤其是‘大英雄’和‘迪阿哥’2个品种表现的抗性较差。(2)随着Cd胁迫浓度的增加,各品种丙二醛（MDA）含量和相对电导率（REC）显著增加,而其保护酶活性不同程度降低。(3)Cd胁迫降低了万寿菊的叶绿素Chl a+b含量和Chl a/b,且抗性差的品种降低的幅度更大;叶片净光合速率（Pⁿ）明显降低,但蒸腾速率(T^r)和气孔导度(G^s)相对降幅较小,对水分利用效率（WUE）影响较小。研究表明,不同品种万寿菊对Cd毒害响应存在明显的差异,且Cd胁迫下生理响应的差异是品种间耐性差异的重要原因之一;抗性差的品种在高浓度Cd胁迫下,光合作用能力降低,抗氧化系统酶的活性受到显著抑制,膜质过氧化加剧,且品种‘珍妮’和‘金门’的抗性强于其他品种,可选择作为Cd污染区栽植植物。     

Chemical forms of aluminum in xylem sap of tea plants (Camellia sinensis L.)

To identify the chemical forms of aluminum (Al) transported from roots to shoots of tea plants (C. sinensis L.), ²⁷Al-nuclear magnetic resonance and ¹⁹F NMR spectroscopy were used to analyze xylem sap.The concentration of Al in collected xylem sap was 0.29 mM, twice as high as that of F. Catechins were not detected in xylem sap. The concentration of malic acid in xylem sap was higher than that of citric acid, whereas the concentration of oxalic acid was negligible.There were two signals in the ²⁷Al NMR spectra of xylem sap, a larger signal at 11 ppm and a smaller one at −1.5 ppm. The former signal was consistent with the peak for an Al-citrate model solution, suggesting that an Al-citrate complex was present in xylem sap. Although the latter signal at −1.5 ppm was thought to indicate the presence of an Al-F complex (at 1.7 ppm) in xylem sap, there was only one signal at −122 ppm in the ¹⁹F NMR spectrum of xylem sap, indicating that the main F complex in xylem sap was F⁻.These results indicate that Al might be translocated as a complex with citrate, while Al-malate, Al-oxalate and Al-F complexes are not major Al complexes in xylem sap of tea plants.     

Distribution of assimilated carbon in plants and rhizosphere soil of basket willow (Salix viminalis L.)

Willow is often used in bio-energy plantations for its potential to function as a renewable energy source, but knowledge about its effect on soil carbon dynamics is limited. Therefore, we investigated the temporal variation in carbon dynamics in willow, focusing on below-ground allocation and sequestration to soil carbon pools. Basket willow plants (Salix viminalis L.) in their second year of growth were grown in pots in a greenhouse. At five times during the plants growth, namely 0, 1, 2, 3 and 4 months after breaking winter dormancy, a subset of the plants were continuously labelled with ¹⁴CO₂ in an ESPAS growth chamber for 28 days. After the labelling, the plants were harvested and separated into leaves, first and second year stems and roots. The soil was analysed for total C and ¹⁴C content as well as soil microbial biomass. Immediately after breaking dormancy, carbon stored in the first year stems was relocated to developing roots and leaves. Almost half the newly assimilated C was used for leaf development the first month of growth, dropping to below 15% in the older plants. Within the second month of growth, secondary growth of the stem became the largest carbon sink in the system, and remained so for the older age classes. Between 31 and 41% of the recovered ¹⁴C was allocated to below-ground pools. While the fraction of assimilated ¹⁴C in roots and root+soil respiration did not vary with plant age, the amount allocated to soil and soil microbial biomass increased in the older plants, indicating an increasing rhizodeposition. The total amount of soil microbial biomass was 30% larger in the oldest age class than in an unplanted control soil. The results demonstrate a close linkage between photosynthesis and below-ground carbon dynamics. Up to 13% of the microbial biomass consisted of carbon assimilated by the willows within the past 4 weeks, up to 11% of the recovered ¹⁴C was found as soil organic matter.     

Physiological changes and growth of micropropagated chile ancho pepper plantlets during acclimatization and post-acclimatization

Little is known about physiological changes that occur with micropropagated chile ancho pepper (Capsicum annuum L. cv. San Luis) plantlets during acclimatization. Plantlets were transferred to ex vitro conditions to study selected physiological changes and growth performance during acclimatization and post-acclimatization. The physiology of the plantlets was characterized by measuring leaf gas exchange and water status. Plant growth was determined by assessing plant height, leaf number, total leaf area, relative growth rate (RGR), and leaf, root, and stem dry matter (DM). Chile pepper plantlets became acclimatized within 6 days after transplantation. During this period, physiological adjustments occurred, which were critical for plantlet survival. After initial ex vitro transplanting, plantlets experienced water deficit [leaf wilting and reduced relative water content (RWC)], which corresponded with reduced stomatal conductance (g _s) and transpiration (E), and an increase in stomatal resistance (r _s). Thus, leaf stomata that developed in vitro were functional ex vitro. Because of this stomatal control, plantlets minimized transplant shock, recovered and survived. Prior to transplanting, plantlets were photomixotrophic, as indicated by low photosynthetic rates (A). During acclimatization, RWC, g _s, E, and A were significantly lower two days after transplanting. However, within 6 days after transplanting, plantlets recovered and became photoautotrophic – attaining high A, g _s, and E. Water use efficiency was initially low during the first days after transplanting, but increased dramatically at the end of the acclimatization period in part due to increased A. The stabilization and improvement of plantlet water status and gas exchange during acclimatization and post-acclimatization closely correlated with increased plantlet growth. This revised version was published online in June 2006 with corrections to the Cover Date.     

Measurement of choline acetyltransferase with [14C]acetate by a cycling procedure

A multiple enzyme and multisubstrate cycling system is described for the radiometric determination of cholineacetyltransferase (ChAT) activity in crude tissue homogenates. The methods employs [¹⁴C]acetate coupled with the enzymes acetate kinase (AK) and phosphotransacetylase (PTA) for the generation of [¹⁴C]acetyl CoA. By recycling it was possible to avoid product inhibition of ChAT by CoA, ATP was maintained constant by rephosphorylation of ADP. Kinetics of the individual enzyme reactions were studied and the parameters obtained were used to select appropriate conditions to maintain linearity of varying amounts ChAT activity over a sixty minute time course. The sensitivity of the method is limited only by the specific activity of commercially available isotope labeled acetate.Special issue dedicated to Dr. O. H. Lowry.     

Field performance and biochemical evaluation of micropropagated mulberry plants

Microclones of different mulberry genotypes were successfully transferred to the field. The same genotypes were raised through conventional methods (cuttings). A comparative study using morphological and biochemical tests of field established micropropagated and cutting derived plants of mulberry genotypes was conducted. Micropropagated mulberry plants showed significant morphogenic vigour when compared to plants raised through cuttings. Biochemical tests of leaves revealed that, there was no significant nutritional difference between micropropagated plants and those originated from cuttings. This revised version was published online in June 2006 with corrections to the Cover Date.     

褶皱臂尾轮虫Brachionus plicatilis摄食小球藻Chlorella sp.的碳同化与碳排放

浮游动物在食物链能量流动与物质循环中发挥着重要作用,能将摄入的浮游植物转化为不同形态的碳,在海洋碳循环中发挥重要作用。应用¹⁴C标记示踪方法,定量分析海洋浮游动物褶皱臂尾轮虫Brachionus plicatilis摄入碳的碳同化与碳排放。喂食不同密度小球藻Chlorella sp.（1×10⁵个/mL、5×10⁵个/mL、1×10⁶个/mL）后,褶皱臂尾轮虫对小球藻碳的同化率（AE）为34%-51%,呈现随饵料密度增加而减小的趋势;未被轮虫同化的碳,主要以溶解有机碳（DOC）的形态排放到水体中,DOC占碳排放的比例为37%-51%,随着饵料密度增加而增加;二氧化碳（CO₂）的比例为15%-40%,随着饵料密度增加而减小;颗粒有机碳（POC）占碳排放的比例较少,为23%-34%,随着饵料密度的增加而增加。此外,分析褶皱臂尾轮虫排放DOC的粒径组成,发现低分子量有机碳（LMW,< 3 kDa）的量大于胶体有机碳（COC,3 kDa-0.22 μm）的量,COC占DOC比例为33%-43%;LMW占DOC比例为57%-67%。本研究结果表明,浮游动物可把相当部分食物中的碳转化为DOC,排放到水体中为细菌所利用,在海洋碳循环中发挥重要作用。