It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column

Benthic macrofaunal populations through their feeding, bioturbation, burrow construction and sediment irrigation activities have profound influences on organic matter inputs to marine sediments (biodeposition) and on the vertical distribution of deposited organic matter within the sediment. These effects in turn influence the rates and pathways of organic matter mineralisation, and element cycles. Similarly, bioturbation, burrow construction and burrow irrigation are major determinants of sediment-water column fluxes of oxygen and nutrients.

In this review, I discuss the influences of the different benthic macrofaunal feeding (functional) groups on mineralisation processes and sediment-water column fluxes of particulate and dissolved nutrients. How these effects influence diagenic processes, the balance between aerobic and anaerobic processes, and the redox status of the surficial sediments. Finally, I discuss some of the limitations of the predominantly laboratory techniques which have been used to study “macrofaunal effects” and how this hinders the inclusion of the effects in quantitative sediment biogeochemical models.     

It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column

Benthic macrofaunal populations through their feeding, bioturbation, burrow construction and sediment irrigation activities have profound influences on organic matter inputs to marine sediments (biodeposition) and on the vertical distribution of deposited organic matter within the sediment. These effects in turn influence the rates and pathways of organic matter mineralisation, and element cycles. Similarly, bioturbation, burrow construction and burrow irrigation are major determinants of sediment-water column fluxes of oxygen and nutrients.

In this review, I discuss the influences of the different benthic macrofaunal feeding (functional) groups on mineralisation processes and sediment-water column fluxes of particulate and dissolved nutrients. How these effects influence diagenic processes, the balance between aerobic and anaerobic processes, and the redox status of the surficial sediments. Finally, I discuss some of the limitations of the predominantly laboratory techniques which have been used to study “macrofaunal effects” and how this hinders the inclusion of the effects in quantitative sediment biogeochemical models.     

Sorption of tetracycline to sediments and soils: assessing the roles of pH,the presence of cadmium and properties of sediments and soils

Batch sorption experiments were conducted to evaluate the sorption behavior of tetracycline (TC, H₃L) on sediments and soils in the presence and absence of cadmium (Cd), as affected by pH and properties of sediments and soils. The results indicated stronger nonlinearity and higher capacity of TC sorption on sediments than on soils. Sorption of TC also strongly depended on environmental factors and sediment/soil properties. Lower pH facilitated TC sorption through a cation exchange mechanism, which also took place at pH values above 5.5, where TC existed as a zwitterion (H₂L⁰) or anions (HL^- and L^2-). When pH was above 7, however, ligand-promoted dissolution of TC might occur due to TC weakening the Al-O bond of aluminum oxide and the Fe-O bond of iron oxide. Natural organic matter (NOM) plays a more important role in TC sorption than cation exchange capacity (CEC) and clay contents. The presence of Cd (II) increased TC sorption on both sediments and soils, which resulted from the decrease of equilibrium solution pH caused by Cd²⁺ exchange with H⁺ ions of sediment/soil surfaces. The increase of TC sorption was also related to the formation of TC-Cd complexes, where Cd²⁺ acted as a bridge between the sediment/soil and TC.     

Biogeochemical zonation of sulfur during the discharge of groundwater to lake in desert plateau (Dakebo Lake,NW China)

As one of the important elements of controlling the redox system within the hyporheic and hypolentic zone, sulfur is involved in a series of complex biogeochemical processes such as carbon cycle, water acidification, formation of iron and manganese minerals, redox processes of trace metal elements and a series of important ecological processes. Previous studies on biogeochemistry of the hyporheic and hypolentic zones mostly concentrated on nutrients of nitrogen and phosphorus, heavy metals and other pollutants. Systematic study of biogeochemical behavior of sulfur and its main controlling factors within the lake hypolentic zone is very urgent and important. In this paper, a typical desert plateau lake, Dakebo Lake in northwestern China, was taken for example within which redox zonation and biogeochemical characteristics of sulfur affected by hydrodynamic conditions were studied based on not only traditional hydrochemical analysis, but also environmental isotope evidence. In the lake hypolentic zone of the study area, due to the different hydrodynamic conditions, vertical profile of sulfur species and environmental parameters differ at the two sites of the lake (western side and center). Reduction of sulfate, deposition and oxidation of sulfide, dissolution and precipitation of sulfur-bearing minerals occurred are responded well to Eh, dissolved oxygen, pH, organic carbon and microorganism according to which the lake hypolentic zone can be divided into reduced zone containing H₂S, reduced zone containing no H₂S, transition zone and oxidized zone. The results of this study provide valuable insights for understanding sulfur conversion processes and sulfur biogeochemical zonation within a lake hypolentic zone in an extreme plateau arid environment and for protecting the lake–wetland ecosystem in arid and semiarid regions.     

Effects of mussel (Perna canaliculus) biodeposit decomposition on benthic respiration and nutrient fluxes

Suspension-feeding bivalves increase the quantity and quality of sedimenting organic matter through the production of faeces and pseudofaeces that are remineralised in coastal sediments and thus increase sediment oxygen demand and nutrient regeneration. Bivalves are intensively cultivated worldwide; however, no bivalve biodeposit decay rates are available to parameterise models describing the environmental effects of bivalve culture. We examined sediment biogeochemical changes as bivalve biodeposits age by incubating coastal sediments to which we added fresh mussel (Perna canaliculus) biodeposits and measured O₂ and nutrient fluxes as well as sediment characteristics over an 11-day period. Biodeposits elevated organic matter, chlorophyll a, phaeophytin a, organic carbon and nitrogen concentrations in the surface sediments. Sediment oxygen consumption (SOC) increased significantly (P=0.016) by ∼1.5 times to 1,010 μmol m⁻² h⁻¹ immediately after biodeposit addition and remained elevated compared to control cores without additions for the incubation period. This increase is in the range of observed in situ oxygen demand enhancements under mussel farms. To calculate a decay rate for biodeposits in sediments we fitted a first-order G model to the observed increase in SOC. The significant model fit (P=0.001, r ²=0.72) generated a decay rate of 0.16 day⁻¹ (P=0.033, SE=0.05) that corresponds to a half-life time of 4.3 day. This decay rate is 1–2 orders of magnitude higher than published decay rates of coastal sediments without organic enrichment but similar to rates of decaying zooplankton faecal pellets. NH₄⁺ release increased rapidly on the day of biodeposit addition (P=0.013) and reached a maximum of 144 μmol m⁻² h⁻¹ after 5 days which was 3.6 times higher compared to control cores. During this period NH₄⁺ release was significantly (P<0.001 to P=0.043) higher in the cores with biodeposit additions than in control cores.     

Sulfur mediated heavy metal biogeochemical cycles in coastal wetlands: From sediments,rhizosphere to vegetation

• In sediments, the transformation of sulfides may lead to the release of heavy metals. • In the rhizosphere, sulfur regulates the uptake of heavy metals by plants. • In plants, sulfur mediates a series of heavy metal tolerance mechanisms. • Explore interactions between sulfur and heavy metals on different scales is needed. The interactions and mechanisms between sulfur and heavy metals are a growing focus of biogeochemical studies in coastal wetlands. These issues underline the fate of heavy metals bound in sediments or released into the system through sediments. Despite the fact that numerous published studies have suggested sulfur has a significant impact on the bioavailability of heavy metals accumulated in coastal wetlands, to date, no review article has systematically summarized those studies, particularly from the perspective of the three major components of wetland ecosystems (sediments, rhizosphere, and vegetation). The present review summarizes the studies published in the past four decades and highlights the major achievements in this field. Research and studies available thus far indicate that under anaerobic conditions, most of the potentially bioavailable heavy metals in coastal wetland sediments are fixed as precipitates, such as metal sulfides. However, fluctuations in physicochemical conditions may affect sulfur cycling, and hence, directly or indirectly lead to the conversion and migration of heavy metals. In the rhizosphere, root activities and microbes together affect the speciation and transformation of sulfur which in turn mediate the migration of heavy metals. As for plant tissues, tolerance to heavy metals is enhanced by sulfur-containing compounds via promoting a series of chelation and detoxification processes. Finally, to further understand the interactions between sulfur and heavy metals in coastal wetlands, some major future research directions are proposed.     

沉积物中汞的生物可利用性研究进展

沉积物是汞在水体中的主要分布相,同时也被认为是许多水生生物摄取汞的主要途径,本文综述了近年来在沉积物中汞的生物可利用性领域所取得的最新研究进展,包括体外消化液萃取法估测沉积物中汞的生物可利用性、可能影响汞生物可利用性的生物地球化学因素(汞在沉积物中的浓度、沉积物的组成、汞与沉积物的接触时间、汞与沉积物的结合形式,以及沉...     

The Influence of Anthropogenic Contamination On Metals Release From Coastal Sediment Suspensions with Aerated Sea Water

The zinc, cadmium, copper and lead release from the uncontaminated and contaminated coastal sediments with aerated sea water was studied. the metals transfer to the dissolved forms was monitored during one-two months by differential pulse anodic voltammetry (DPASV). the sediments with different initial degree of contamination were sampled in the Peter the Great Bay, Sea of Japan around Vladivostok-the biggest city in Russian Far East. Sediment contamination by metals led to increased release to solution of zinc due to sulphide oxidation and cadmium on account of organic matter decomposition. the copper behaviour was complicated by strong binding with organic matter and enhancements of copper release can only be seen in sediment with a low organic matter content. Significant lead transfer to dissolved forms was not observed regardless of sediment contamination. the temperature affected the release of cadmium and copper through enhanced organic matter destruction. the dissolved metal fluxes from the sediment transformation are compared with metal fluxes towards bottom. Such comparisons show that second contamination by dissolved metals of the studied coastal environment may be important for cadmium only.     

The Influence of Anthropogenic Contamination On Metals Release From Coastal Sediment Suspensions with Aerated Sea Water

The zinc, cadmium, copper and lead release from the uncontaminated and contaminated coastal sediments with aerated sea water was studied. the metals transfer to the dissolved forms was monitored during one-two months by differential pulse anodic voltammetry (DPASV). the sediments with different initial degree of contamination were sampled in the Peter the Great Bay, Sea of Japan around Vladivostok-the biggest city in Russian Far East. Sediment contamination by metals led to increased release to solution of zinc due to sulphide oxidation and cadmium on account of organic matter decomposition. the copper behaviour was complicated by strong binding with organic matter and enhancements of copper release can only be seen in sediment with a low organic matter content. Significant lead transfer to dissolved forms was not observed regardless of sediment contamination. the temperature affected the release of cadmium and copper through enhanced organic matter destruction. the dissolved metal fluxes from the sediment transformation are compared with metal fluxes towards bottom. Such comparisons show that second contamination by dissolved metals of the studied coastal environment may be important for cadmium only.     

Long-term geochemical evolution of acidic mine wastes under anaerobic conditions

A nearly 5-year anaerobic incubation experiment was conducted to observe the geochemical evolution of an acidic mine waste. Long-term storage of the mine waste under strict anaerobic conditions caused marked increase in aqueous sulfur, while aqueous iron showed no remarkable change. Co-existing oxidation and reduction of elemental sulfur appeared to play a central role in controlling the evolutionary trends of aqueous sulfur and iron. Addition of organic matter increased the aqueous Fe concentration, possibly due to enhanced iron mobilization by microbial iron reduction and increased iron solubility by forming organically complexed Fe species. Further addition of CaCO₃ resulted in immobilization of aqueous iron and sulfur due to elevated pH and gypsum formation. The chemical behaviors of environmentally significant metals were markedly affected by the added organic matter; Al, Cr, Cu, Ni and Zn tended to be immobilized probably due to elevated pH and complexation with insoluble organic molecules, while As and Pb tended to be mobilized. Jarosite exhibited high stability after nearly 5 years of anaerobic incubation and even under circumneutral pH conditions. Long-term weathering of aluminosilicate through acid attack raised pH, while continuous reaction between the added CaCO₃ and mine waste-borne stored acid decreased pH.     

Experimental degradation of oil in permeable sand from the Gulf of Aqaba,Red Sea

Degradation rates of light and heavy oil in permeable carbonate sands from the Gulf of Aqaba were investigated to evaluate the ability of sediments to degrade oil compounds. Silicate sands that are less permeable and different properties from carbonate sands were used for comparison. Estimates of oil degradation rates were based on oxygen consumption rates, calculated by incubating natural carbonate sands with oil. The degradation rates of light oil were twofold higher than those of heavy oil, which may be attributed to the presence of a higher carbon number in heavy oil compared with light oil. Degradation rates of light oil in carbonate sands were twofold higher than in silicate sands. Oil degradation rates calculated using the bottle incubation technique were three- to fourfold higher than rates from chamber incubations, indicating the importance of adequate mixing between oil particles and sediments during degradation processes. This study suggests that permeable sands, through their chemical and physical properties, increase oil biodegradation rates by enhancing flow through sediment particles and positively impacting bioturbation processes.     

Feeding and bacteriolytic responses of the deposit-feeder Abarenicola pacifica (Polychaeta: Arenicolidae) to changes in temperature and sediment food concentration

Deposit-feeders can respond to seasonal fluctuations in food concentration both functionally (e.g. by adjusting feeding rates) and physiologically (e.g. by changing the concentration of bacteriolytic agents in gut fluids). Laboratory feeding experiments were carried out (11 to 21 July 1997) with the arenicolid polychaete worm Abarenicola pacifica (Healy and Wells). Objectives were to test for separate and interactive effects of sediment food concentration and temperature (6, 11, and 16 °C) on deposit-feeder functional (feeding rates) and physiological (bacteriolytic activity of gut fluids) responses. Food concentration was varied experimentally using sieved (1 mm) natural sediments (Md φ=2.00; 0.6% organic) mixed with combusted (500 °C, 8 h) sediments for final concentrations of 25, 50, and 100% natural sediment. Sediment food quality was measured as: (1) bioavailable amino acids (EHAA), (2) chlorophyll a (chl a), and (3) bacterial abundance. Feeding rates were inferred from egestion rates (ER, g h⁻¹) and analyzed with respect to worm size. Bacteriolytic activity of midgut fluids was assayed turbidimetrically against two bacterial isolates, after worms had fed on experimental sediments for 15 d. Temperature and food concentration both significantly affected feeding rates, with maxima occurring at 50 and 100% natural sediment mixtures, and at high (16 °C) temperature. ER was positively, but not significantly correlated with EHAA and chl a; a positive, significant correlation was detected between ER and sediment bacterial abundance. Overall, functional responses agreed with earlier compensatory intake models for deposit-feeders. However, the size and direction of these responses was temperature-sensitive, suggesting that these models need to be adjusted for changes in absorption rates. No effects of ambient temperature or food concentration on bacteriolytic rates were observed, possibly due to compensatory mechanisms or the presence of multiple bacteriolytic agents in gut fluids. Received: 28 June 1999 / Accepted: 14 March 2000     

曝气复氧对底泥氮素生物地球化学循环影响的作用机制研究

通过分析底泥氮污染物释放规律和转化过程,以及底泥生境、氮形态变化和氮循环功能微生物群落结构变化的规律,探讨了不同曝气复氧条件影响底泥氮生物地球化学循环的生物代谢、物理化学联合作用的机制。结果表明：曝气复氧对底泥中氮的生物地球化学循环影响是一个包括微生物代谢作用和物理化学作用的复杂联合作用过程。水体好氧环境的改变主要引起参与底泥氮循环的硝化、亚硝化和反硝化功能菌群群落结构的演变,对异养菌和氨化菌的影响不大,证明环境好氧条件的改变对底泥有机质生物分解产生氨氮的微生物代谢过程影响不大,主要对底泥释放的氨氮硝化、反硝化等生物转化过程产生大的影响。不同溶解氧条件下,底泥释放的氮素在微生物作用下主要以NH4＋-N和NO3--N的形式进入试验体系,并在特定的氧化还原电位（临界值-200 mV）和pH（临界值6.70）条件下通过物理化学作用在底泥中以离子交换态氮（IEF-N）、碳酸盐结合态氮（CF-N）、铁锰氧化态氮（IMOF-N）及有机态和硫化物结合态氮（OSF-N）等不同形态氮相互转化,同时,在氮的转化和循环过程中部分输入上覆水体。在低溶解氧组实验条件下[ρ（DO）〈0.5 mg.L-1],底泥向水体输出氮总量为底泥可转化态氮的19.7%,主要为氨氮,最大释放速率达到289.13 mg.m-2.d-1,释放的质量浓度可达到18.8 mg.L-1;好氧条件下（DO饱和）,底泥向水体输出氮总量为底泥可转化态氮的1.8%;好氧-缺氧条件下为11.7%,主要以N2的形式释出系统。     

Laboratory evaluation of kinetic parameters for lake sediment denitrification models

Laboratory experiments using the acetylene blockage technique showed that denitrification rates in sediments from Lake Okeechobee, Florida, are described by Michaelis-Menten nitrate dependence and Arrhenius temperature dependence (E_act = 15.5 kcal, but do not appear to be affected by seasonal differences in the supply of carbon substrates in this eutrophic lake. Peak nitrate concentrations in the sediments are in the range of the half-saturation constant, and thus, for a given temperature, k_d varies by a factor of approximately two over the typical range of nitrate concentrations. Most diagenetic sediment denitrification models employ a single-valued rate constant k_d, to describe nitrate dependence. However, the use of a constant k_d for eutrophic lake sediments may result in a significantly erroneous estimate of annual denitrification loss.     

Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3–28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2–1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

Impact of the polychaete Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon fluoranthene

Polychaetes belonging to the genus Capitella are often present in high numbers in organic-rich sediments polluted with, e.g., oil components, and Capitella spp. may have a great impact on the biogeochemistry of these sediments. We examined the influence of Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon, fluoranthene. Capitella sp. I were added to microcosms (10 000 ind m⁻²) and the impact of a pulse-sedimentation of fluoranthene-contaminated sediment (3 mm layer) was studied for a period of 12 d after sedimentation. The sediment oxygen uptake and total sediment metabolism (TCO₂ production) increased in cores with worms (71 to 131%), whereas the anaerobic activity, measured as sulfate reduction rate 12 d after sedimentation, was lower compared to cores without worms. The effect of fluoranthene on sulfate reduction was most pronounced in the presence of worms, with a 34% reduction versus 16% in cores without worms. The reduced sulfur pools in cores with worms were smaller than in cores without worms, suggesting that the reduced anaerobic activity was caused by increased oxidation of the sediment, which may favor O₂ and other electron-acceptors (e.g. NO₃ ⁻, Fe³⁺, Mn⁴⁺) in organic matter decomposition. The sediment oxygen uptake and TCO₂ production did not show significant changes due to fluoranthene treatment, indicating that these parameters were either less sensitive to fluoranthene stress or recovered more rapidly (i.e. within 48 h) than sulfate reduction rates. Bioturbation by Capitella sp. I altered the depth profile of fluoranthene such that fluoranthene was found in deeper sediment layers (down to 2 cm) where diffusional loss and microbial breakdown probably are reduced relative to surface layers. In cores without worms, fluoranthene was found down to 1 cm, with 75% remaining in the upper 5 mm. Received: 5 December 1996 / Accepted: 11 February 1997     

Seasonal variability of intertidal bacterial metabolism and growth efficiency in an exposed sandy beach during low tide

Benthic gross primary production (GPP) and community respiration (BCR) were regularly measured on sandy beach sediment during low tide in a megatidal ecosystem. These measurements were assessed during 2 years in situ within a benthic chamber simultaneously with bacterial production (BP). Results suggested that community respiration in Wimereux sandy beach sediments was probably dominated by bacterial respiration and that the estimated benthic bacterial growth efficiency (BGE) was highly variable. Although temperature significantly controlled both BP and BCR during low tide at the annual scale, the temperature effect on BGE was not significant. The instability and sediment erosion caused by the wave action and the tidal cycle were thus thought to influence the BGE. Since the sampling site is regularly affected by Phaeocystis foam deposit (enhancing the BCR and decreasing the BGE), and since GPP rates were highly variable and supported by high assimilation numbers (i.e., >1 mgC mgChla ⁻¹ h⁻¹), phytoplankton organic matter deposit following the immersion of the study site was thought to explain the BGE variability.     

Effect of monsoonal climate on sulfate reduction in coastal sediments of the central Great Barrier Reef lagoon

Rates of sulfate reduction were measured during the spring dry and summer wet seasons of 1992 in shallow coastal sediments of the central Great Barrier Reef lagoon. In the dry season, sulfate reduction rates, integrated to 18–20 cm depth, ranged from 23.8 to 30.8 mmol S m^-2 d^-1. In the wet season, heavy monsoonal rains and wind-induced waves caused severe disturbance of sediments leading to less anoxic conditions, and to a 2- to 3-fold decline in rates of sulfate reduction (10.2 to 12.8 mmol S m^-2 d^-1). The percentage of reduced ³⁵S recovered as acid-volatile sulfide (% AVS_red) ranged from 15.8 to 44.9% in spring, and was significantly reduced at each station in summer (range 14.5 to 31.4%). An analysis of variance indicated that seasonality accounted for 31.5% of the total variance in total rates of sulfate reduction, with only 7.5% of the variance accounted for by depth into the sediment; there were no station differences. In both seasons, there were no clear patterns in the proportion of radiolabel incorporated into the AVS and chromium-reducible sulfur (CRS) pools with depth into the sediment. These results are contrary to the seasonal patterns and pathways of sulfate reduction in temperate coastal sediments.     

Chromium in intertidal sediments of the Clyde,UK: potential for remobilisation and bioaccumulation

Detailed surveys of intertidal sediments have been performed along the north and south shores of the Inner Clyde estuary, UK. Surface sediment data reveal significant spatial variation in Cr content and an association with major sediment characteristics and location within the estuary. Depth variation for Cr and other heavy metals cannot be explained by variation in major geochemical controls such as grain size and organic matter and highlights the impact of historical contamination on sediment quality. These elevated levels at depth may still have environmental impact through redox-reactivity, in association with iron and manganese. Sequential extraction of sediments and pore water analysis of Cr(VI) and Cr(III) provide detailed information on release potential from the sediments. The implication of Cr mobility for biota in the estuary has been assessed by the analysis of a common marine bivalve, Mytilus edulis (Blue Mussel) and a burrowing polychaete, Nereis diversicolor (rag worm) from a number of survey sites. Bioconcentration factors for Mytilus indicate that the weakly held portion of sediment Cr is available for uptake and in the case of Nereis, bioaccumulation appears to be inhibited by sediment organic matter.     

黄河上游沉积物对磷酸盐的吸附动力学研究

研究了黄河上游10个不同表层沉积物在黄河水体中对磷酸盐（P）的吸附动力学及其影响因素和吸附机理。结果表明：不同黄河沉积物对P的吸附能力各不相同,但吸附量随时间的变化具有相同的趋势,吸附速率均在前8 h内较快,以后逐渐趋缓,在48 h时基本达到吸附平衡。不同黄河沉积物对P的吸附量均随P初始质量浓度的增加而增大,随沉积物质量浓度增大而减小,且也受水体pH值的影响,在pH为6.0~9.0范围内吸附量比较大。不同沉积物在不同P起始质量浓度下对P的吸附动力学均符合Lagergren二级吸附动力学模型及Weber–Morris扩散方程,求得二级吸附速率常数和扩散速率常数分别在10.85~229.29 g.mg-1.h-1和0.7×10-3~5.2×10-3 mg.g-1？h-1/2）之间,吸附过程由P在沉积物内的扩散控制。