Adenosine 5'-triphosphate flux through the north inlet marsh system

The distribution, fluctuation, and short-term transport of total microbial biomass (measured as adenosine 5'-triphosphate [ATP]) was investigated in a large salt marsh creek. Hourly samples were collected synoptically for 25 h from 10 boats positioned across the 320-m width of the creek. Samples were collected from three depths ranging from 0.2 to 8.0 m. Hourly data obtained from each station were graphed, plotting depth against ATP. Subsequently, interpolated ATP values were generated for every one-tenth depth from the surface to the bottom with the use of an 11-point proportional divider. A total of 2,750 values were generated, and a mean value of 0.865 mg of ATP per m was determined. Maximum levels of ATP were found at high tide and minimal values were found at low tide. The distribution of ATP concentrations was found to be complex, with no suggestion of vertical stratification; however, horizontal divisions were apparent. ATP values corrected for direction of flow or velocity indicated two ebb-directed channels; however, when considered in total, there was a net import of ATP through the interface. The total import of ATP for this 25-h sampling period was calculated to be 3.58 kg, corresponding to a net transport of 39.8 mg of ATP per s through the cross section. Results suggest that detailed characterization of a creek transect in terms of ATP or any similar parameter requires the simultaneous measurements of both the concentration of the parameter in question and the velocity at the time and point from which the sample was taken.     

The transport of microbial biomass and suspended material in a high-marsh creek

The transport of microbial biomass and suspended material in a high-marsh creek was investigated during four 40-h tidal studies throughout the year. Although considerable differences were noted between successive tidal cycles, overall the creek was found to be an exporting system and transported a mean concentration of ATP (-33g), chlorophyll a (-66g), particulate organic carbon (-31kg), total suspended material (-344kg), and fixed suspended material (-195kg) during each tidal cycle. This net outward flux of materials was associated with a net flow of water out of creek, while the net import of aerobic, heterotrophic bacteria (43 x 10(12)) and volatile suspended material (238kg) was generally due to higher mean concentrations of these materials per unit volume of water during the flooding tide. Also the latter generally were associated with increased amounts of suspended material suggesting an association between bacteria and suspended matter.     

Transport of microbial biomass through the North Inlet ecosystem

Tidal fluctuations and transports of total microbial biomass (measured as adenosine triphosphate [ATP]) were investigated at three marsh creeks comprising the major transfer points between the North Inlet marsh and the adjoining aquatic ecosystems. Two creeks, Town Creek and North Jones Creek, form the inlet mouth and are the only marsh-ocean exchange points. The third creek, South Jones Creek, connects to a brackish water embayment. The creeks were simultaneously sampled every 1.5 hours for 50 consecutive hours during neap tides (four tidal cycles) and 50 consecutive hours during spring tides of each season. At the inlet, ATP concentration fluctuated in phase with the tide during winter and fall and out of phase with the tide during the spring. Fluctuation patterns at South Jones Creek were irregular. The highest ATP concentrations were during the spring (mean=2.17 mg of ATP per m³) and the lowest concentrations were during the winter (mean=0.65 mg of ATP per m³). Net transports of ATP varied from tidal cycle to tidal cycle with regard to direction of transport (import or export) and magnitude. Net transports were small compared to large instantaneous transports and only 4 of 22 determinations of net transport were different from zero transport.     

Tidal and diurnal variations in larval fish abundance in an estuarine inlet in Ariake Bay,Japan: implication for selective tidal stream transport

We have conducted a preliminary study of tidal and diurnal variations in the distribution of dominant larval and juvenile fishes in the Chikugo River inlet (Ariake Bay, Kyushu, Japan) to determine whether selective tidal stream transport (STST) occurs. Larval and juvenile fish were collected from the mesohaline zone of the Chikugo River inlet during spring 2002. Temperature, salinity, depth, and current velocity were measured. Larval and juvenile abundance were compared among four tidal conditions, flooding tide, high tide, ebbing tide, and low tide, and between day and night. A total of 12 families, 15 species, and 5,577 individuals were collected. Temperature did not vary significantly with tidal conditions whereas salinity, depth, and current velocity varied significantly. Salinity also was correlated significantly and positively with depth. The abundance of most of the fishes was correlated positively and significantly with salinity and depth. Lateolabrax japonicus, Trachidermus fasciatus, Acanthogobius hasta, and other gobiid larvae (Gobiidae spp.) were significantly more abundant during high tide; in contrast, Coilia nasus and Neosalanx reganius were most abundant during low tide. The abundance of most of the fishes was higher during high tides at night than during the day, indicating the existence of STST, which may be strategically associated with ascending progress to upstream nursery areas.     

Flux of decapod larvae and juveniles at a station in the lower Canal de Mira (Ria de Aveiro,Portugal) during one lunar month

Summary

Emigration and immigration of decapod larvae from estuaries depend on timing of larvae occurrence in the water column relative to the tidal, tidal amplitude and day cycles. The phase relation of these natural cycles varies with tidal regime and geographically, resulting in different time-patterns of hatching of first stage larvae and of presence of late stage larvae in the water column. Vertical migration behaviour according to phase of tide also controls transport inside estuaries. These mechanisms were investigated in a field study conducted on the northwest coast of Portugal where neap ebb tides occur during the night around the quarters of the moon. Flux of decapod larvae through one sampling station was measured during one lunar month at the Canal de Mira (Ria de Aveiro) in the spring of 1990. The sampling programme was comprised of a set of 25-h fixed station studies, separated by 25-h intervals during which no sampling took place. Plankton samples were collected with a pump every hour at three depths. Current velocity and direction at the standard depths, as well as height of the water column, were also measured every hour. Hourly instantaneous flux of larvae through a 1-m-wide vertical section of the Canal de Mira was calculated for the most abundant forms. A total number of 13 combinations of species and larval stages were analyzed, belonging to the families Atelecyclidae, Pirimelidae, Portunidae, Pilumnidae, Grapsidae, Palaemonidae, Crangonidae and Thalassinidae. Patterns of net larval flux along the lunar month could be grouped into three types. Type 1 includes first zoeas that were consistently exported to the sea. Type 2 comprises late zoeas, megalops and juveniles that were consistently imported into the estuary. First zoeas that were imported during some of the 25-h studies but were exported during the others were included in Type 3; in species of this type import periods appeared to alternate with export periods according to lunar phase. Flux of Type 1 larvae followed a semi-lunar pattern. Release activity of Type 1 zoeas took place during the night and started during neap tides around the quarters of the moon, but maximum releases occurred 3–4 h after high tide of average amplitude tides, 3–4 days after the quadratures. These observations agree with the hypothesis that hatching is timed to occur on ebb tides of the largest possible amplitude so that larvae are easily dispersed from areas with a high density of predator fishes. However, based on other observations on the Portuguese coast, it cannot be ruled out that hatching might depend on a minimum number of hours of darkness experienced by the females. Larvae included in Type 2 comprise forms that may have been retained inside the estuary for the entire larval phase, as well as one form that was imported from shelf waters. No semi-lunar pattern of import was detected in this last form. Fluctuations of net flux observed in Type 3 larvae, as well in other forms that were not included in any of the types, were more difficult to explain. These larvae were first zoeas of species belonging to different taxonomic, morphological and ecological groups and may show a diversity of adaptations to the way of life of the adults. Imports and exports of larvae depended not only on time-patterns of abundance, but also on time-patterns of larval vertical distribution. As a general rule, larval stages showed patterns of depth distribution that were consistent with vertical migration rhythmic behaviours synchronized with the tidal cycle. Though the effect was not always statistically significant, first-stage larvae were closer to the surface during ebb, especially during the night, enhancing seaward transport. On the contrary, later zoeal stages, megalops and juveniles were usually closer to the surface during flood, suggesting migration to the water column during this phase of the tide and landward transport.     

Length‐weight relationships of 4 fish species from a seagrass Zostera marina bed in Koje Bay,Korea

Length‐weight relationships of 4 fish species collected from a seagrass Zostera marina bed in Koje Bay on the southern coast of Korea were analyzed. The fish samples were captured using a seine net (mesh size 6 mm) at water depth of approximately 1.5 meters during low tide. Fishes were collected 8 times from summer 2016 to spring 2017. The regression parameter b varied between 2.47 for Favonigobius gymnauchen and 3.03 for Ditrema temminckii temminckii.     

The relation of sodium and potassium ion transport to the respiration and adenine nucleotide content of liver slices treated with inhibitors of respiration

1. The dependence of the net transport of Na(+) and K(+) by rat liver on the respiration has been determined by incubating slices in the presence of varying concentrations of respiratory inhibitors. 2. Neither the rate of net transport nor the total amount of each ion transported was inhibited unless the rate of endogenous respiration was decreased below a critical value of about 330mmol of O(2)/h per kg of protein (i.e. 50% of the total endogenous respiration). 3. The uninhibited rate of respiration could be varied over a twofold range (380-770mmol of O(2)/h per kg of protein) by the use of different substrates, but the critical value for the onset of transport inhibition was quite constant (290-360mmol/h per kg of protein) under these different conditions. 4. Slices incubated at 38 degrees C without inhibitors showed an increase of their ATP content and the concentration ratio ATP/ADP. The final ATP content and concentration ratio, ATP/ADP, of slices treated with different concentrations of inhibitors were closely related to the rate of respiration. 5. The increased ATP content of the control slices during incubation was equal to the increase of total adenine nucleotides. At increasing degrees of respiratory inhibition the relative contributions of ADP and AMP to the total adenine nucleotide content increased. 6. The critical rate of respiration for the onset of inhibition of ion transport and the corresponding contents of adenine nucleotides provide estimates of the maximal values of certain parameters of energy metabolism required for the support of alkali-cation transport in the liver slices.     

The contribution of bacteria to the total adenosine triphosphate extracted from the microbiota in the water of a salt-marsh creek

The contribution of bacteria to the total adenosine 5-triphosphate (ATP) measured in estuarine samples was investigated. A portion of the bacterial population present in samples was separated from other biological materials by filtration through a 1.0-μm filter (Nucleopore). The quantification of ATP and the enumeration of bacteria (by epifluorescence microscopy) was performed on filtered and unfiltered portions of each sample as well as on cultured organisms. The values of ATP per cell averaged 1.48 × 10⁻¹⁵ g/cell from laboratory cultures and generally two orders-of-magnitude lower in bacteria from the natural system with values ranging from 0.24 to 28.8 × 10⁻¹⁷ g/cell. In bacteria collected from the marsh waters, the higher values of ATP per cell were observed in samples collected during the warmer months. When samples were obtained from areas of the marsh influenced by oceanic waters, the highest values of ATP per cell were observed at low tide. The nucleotide levels in bacterial cells in the high marsh were not markedly influenced by the tides. Bacterial ATP contributed from < 1–83% of the ATP measured in the total microbiota; the average content was 25% of the total. Generally, waters from the high marsh i.e., water influenced strongly by sediment microbes, contained a greater percentage of bacterial ATP.     

Energetics of Anaerobic Sodium Transport by the Fresh Water Turtle Bladder

Certain of the metabolic events associated with anaerobic sodium transport by the isolated bladder of the fresh water turtle have been investigated. The data suggest that energy for this transport arises from glycolysis and that endogenous glycogen was the major and perhaps the sole source of substrate. The rate of anaerobic glycolysis, as determined by lactate formation, correlates well with the rate as determined by glycogen utilization. Using lactate formation as the index of anaerobic glycolysis, a linear relationship was observed between glycolysis and net anaerobic sodium transport. In the absence of sodium transport, glycolysis decreased by approximately 45 per cent. Tissue ATP concentrations were maintained at about the same level under anaerobic as under aerobic conditions. Finally if it is assumed that in the conversion of glycogen to lactate anaerobically, 3 moles of ATP are generated per mole of glucose residue, an average of over 15 equivalents of sodium were transported for every mole of ATP generated.     

养殖排放农药和抗生素在红树林区中残留的初步研究

广西北海大冠沙红树林区是养殖废水的排放地,该文报道2008年10月农药和抗生素在该林区红树林滩、沙滩和潮沟三类生境的残留水、沉积物和底栖动物中的分布与含量。通过对103种农药和22种抗生素检测,发现该林区有16种农药,4种抗生素,其中甲氰菊酯、仲丁威和醚菌酯是主要的污染农药。在残留水中检出12种农药,它们在红树林滩、沙滩和潮沟中的总浓度分别是42.70、29.63和27.41ng/L。在沉积物中检出10种农药,它们在红树林滩、沙滩和潮沟中的总浓度分别是7.90、6.70和7.29μg/kg.DW。在底栖动物体内检出4种农药和3种抗生素。分析表明,红树林滩(残留水和沉积物)比底栖动物更易受到养殖废水的污染,底栖动物对抗生素的富集能力似乎高于对农药的富集能力,生境间污染相似度高于样品间的污染相似度。     

Ontogenetic changes in diel activity,food habits and spatial distribution of juvenile and adult creek chub,Semotilus atromaculatus

Synopsis Juvenile creek chub, Semotilus atromaculatus, differed ecologically from adults. They were essentially diurnal, fed on small prey (adult diptera and aquatic adult coleoptera) and were found mainly in shallow littoral water (0.8 m depth) of a Québec oligotrophic lake. In contrast, adults were principally nocturnal, fed on larger prey (Gammaridae, diptera larvae, diptera pupae and ephemeroptera larvae) and were found in deeper water (1.2, 2.0 and 2.9 m depth). The overlap in diet between juvenile and adult fish was 10 percent. We found that predation by brook charr, Salvelinus fontinalis, upon creek chub was low, as only 5 chart stomachs of the 302 examined contained a total of 16 juvenile chub and none contained adult chub. Laboratory observations revealed that under low light intensity (0.17 lux) adult creek chub fed as efficiently as under high light intensity (22 lux). Several possible hypotheses to explain the evolution of the two different strategies exhibed by adult and juvenile creek chub are discussed.     

Association of Large Juvenile Red Drum, Sciaenops ocellatus, with an Estuarine Creek on the Atlantic Coast of Florida

Seasonal abundance, size distribution, year-class presence, residence duration, and migrational patterns of red drum, Sciaenops ocellatus, in an estuarine marsh creek in the northern Indian River Lagoon, Florida, were investigated during a study in which gill net samples were collected monthly from August 1991 to March 1996. A total of 282 large juvenile red drum were collected, of which 161 were tagged and released and 68 were sacrificed for age determination. Although red drum were collected in the creek throughout the year, abundance levels were negatively correlated with water temperatures and reached maximum levels during the winter months. Significant correlations between fish abundance and salinity or dissolved oxygen levels were not detected. The majority (90%) of the fish collected were large juveniles (260–450mm standard length) that were estimated to be from 10 to 26 months old. The oldest red drum we examined from the creek was estimated to be 37 months old. Tag-recapture data indicated that some fish repeatedly used or were associated with the creek for periods of up to 18 months after release. Estuarine creeks in this region provide exploitable habitat for large juvenile red drum (相似文献   

环境因子对东海区帆式张网主要渔获物渔获量影响

帆式张网渔业是东海区重要的渔业捕捞方式之一．利用广义加性模型(GAMs)定量研究了海表水温、渔场水深、渔场位置和潮汐(农历日期)等环境因子对东海区帆式张网主要渔获物渔获量变动的影响．结果表明,海表水温和渔场水深是影响带鱼、小黄鱼和鲳鱼单位努力渔获量(CPUE)的主要因子,且各因子的影响程度不尽相同,多呈非线性相关关系．从模型的拟合结果来看,海表水温对带鱼的影响最大,其次是水深,渔场位置和潮汐的影响很小．渔场水深对小黄鱼的影响最大,海表水温、渔场位置和潮汐的影响比较接近．渔场水深和海表水温是影响鲳鱼单位努力渔获量的主要环境因子,影响程度比较接近．     

Zonal distribution and population biology of Ilyoplax frater (Brachyura: Ocypodoidea: Dotillidae) in a coastal mudflat of Pakistan

Zonal distribution and population biology of Ilyoplax frater were studied in a mangrove mudflat area of Pakistan. The crabs were collected from Korangi creek through transect and quadrat method from low tide level to high tide level. Two transects were delimited in a mangrove area of Korangi creek (24°79'N/ 67°20'E). On each transect, three 0.25 m quadrats were sampled at three tidal levels on a monthly basis during low tide periods from March 2001 to February 2002. A total of 1124 crabs were obtained, of w...     

The turbidity maximum in the Tamar estuary

Many estuaries of medium to high tidal range exhibit an accumulation of fine cohesive material in their upper reaches in the region of the limit of saline intrusion. Much, or all, of this material is suspended each tidal cycle and the entire region undergoes a seasonal variation which appears to depend on fluvial input. Two factors which are throught to influence the formation and maintenance of turbidity maxima are the differing magnitudes of the bed shear stress (τ₀) on flood and ebb tides and the large vertical density gradient which developes on the ebb tide. Crucial to the importance of the first factor is that τ₀ exceeds a critical value, at which erosion occurs, for a greater period on the flood than on the ebb. The effect of the density gradient is that upward propagation of bed generated eddies is inhibited and the sediment is not transported into the upper part of the flow where it will be most effectively transported. It is not clear which, if either, of these mechanisms is dominant. Data consisting of vertical profiles of velocity, salinity and suspended solids were collected at four stations in the Tamar estuary during a high range tidal cycle. One station, at which the depth mean salinity (S _d ) varied from 0.0 to ∼ 12.0‰, was occupied permanently. The other stations were occupied such that data were collected asS _d varied in the range 0.0 to ∼ 4.0‰. In this way each station was occupied for a period of time on the ebb and flood tide. Observations show that during the early ebb, when the flow is relatively deep and slow, stratification persist untilS _d ∼ 0.0‰ and that no significant transport occurs while the flow is saline but that there is a rapid increase in suspended solids concentrations after this time. During the later ebb the shallower faster flow allowed the density gradient to be erode and significant transport was observed atS _d ∼ 5.0‰. On the flood tide the flow in the low salinity region is well-mixed troughout. Computation of the fluxes and total transport per unit breath of estuary show that on the ebb tide the quantity of solid material being transported by the low salinity (0–3‰) region remains nearly constant as this region of the flow is advebted seaward. On the flood tide, however, as the same region is advected landward the quantity of material being transported increases. It is concluded that in the Tamar estuary the early ebb tide stratification contributed to the formation and maintenance of a turbidity maximum which is strongly associated with the low salinity region of the flow. It is also speculated that the differences in the ebb and flood tide transport are caused by differences in the availability of mobile material on the bed at different stages of the tidal cycle.     

The broad range di- and tri-nucleotide exchanger SLC35B1 displays asymmetrical affinities for ATP transport across the ER membrane

In eukaryotic cells, uptake of cytosolic ATP into the endoplasmic reticulum (ER) lumen is critical for the proper functioning of chaperone proteins. The human transport protein SLC35B1 was recently postulated to mediate ATP/ADP exchange in the ER; however, the underlying molecular mechanisms mediating ATP uptake are not completely understood. Here, we extensively characterized the transport kinetics of human SLC35B1 expressed in yeast that was purified and reconstituted into liposomes. Using [α³²P]ATP uptake assays, we tested the nucleotide concentration dependence of ATP/ADP exchange activity on both sides of the membrane. We found that the apparent affinities of SLC35B1 for ATP/ADP on the internal face were approximately 13 times higher than those on the external side. Because SLC35B1-containing liposomes were preferentially inside-out oriented, these results suggest a low-affinity external site and a high-affinity internal site in the ER. Three different experimental approaches indicated that ATP/ADP exchange by SLC35B1 was not strict, and that other di- and tri-nucleotides could act as suitable counter-substrates for ATP, although mononucleotides and nucleotide sugars were not transported. Finally, bioinformatic analysis and site-directed mutagenesis identified that conserved residues K117 and K120 from transmembrane helix 4 and K277 from transmembrane helix 9 play critical roles in transport. The fact that SLC35B1 can promote ATP transport in exchange for ADP or UDP suggest a more direct coupling between ATP import requirements and the need for eliminating ADP and UDP, which are generated as side products of reactions taking place in the ER-lumen.     

Role of climate and agricultural practice in determining matter discharge into large,shallow Lake Võrtsjärv,Estonia

Understanding the dynamics of fine sediment transport across the upper intertidal zone is critical in managing the erosion and accretion of intertidal areas, and in managed realignment/estuarine habitat recreation strategies. This paper examines the transfer of sediments between salt marsh and mudflat environments in two contrasting macrotidal estuaries: the Seine (France) and the Medway (UK), using data collected during two joint field seasons undertaken by the Anglo-French RIMEW project (Rives-Manche Estuary Watch). High-resolution ADCP, Altimeter, OBS and ASM measurements from mudflat and marsh surface environments have been combined with sediment trap data to examine short-term sediment transport processes under spring tide and storm flow conditions. In addition, the longer-term accumulation of sediment in each salt marsh system has been examined via radiometric dating of sediment cores. In the Seine, rapid sediment accumulation and expansion of salt marsh areas, and subsequent loss of open intertidal mudflats, is a major problem, and the data collected here indicate a distinct net landward flux of sediments into the marsh interior. Suspended sediment fluxes are much higher than in the Medway estuary (averaging 0.09 g/m³/s), and vertical accumulation rates at the salt marsh/mudflat boundary exceed 3 cm/y. Suspended sediment data collected during storm surge conditions indicate that significant in-wash of fine sediments into the marsh interior can occur during (and following) these high-magnitude events. In contrast to the Seine, the Medway is undergoing erosion and general loss of salt marsh areas. Suspended sediment fluxes are of the order of 0.03 g/m³/s, and the marsh system here has much lower rates of vertical accretion (sediment accumulation rates are ca. 4 mm/y). Current velocity data for the Medway site indicate higher velocities on the ebb tide than occur on the flood tide, which may be sufficient to remobilise sediments deposited on the previous tide and so force net removal of material from the marsh.     

Behavior of the groundwater in a riverine-type mangrove forest

The behavior of groundwater and physical properties of bottom sediment in a riverine-type mangrove forest which is composed of a tidal creek and fringing mangrove swamps were investigated through field observations at Iriomote Island, Japan. After the tidal water ebbed from the swamp surface to the creek, groundwater levels at swamp sites near the creek fell by up to 15 cm by the next flood tide, although the fall was negligible at sites far from the creek and at the open coast outside the mangrove forest. The amount of groundwater discharged to the creek from the swamp depended strongly both on the tidal range and the presence of the steep bank which separates the tidal creek from the fringing mangrove swamp. Based on the fall of groundwater level, the bulk hydraulic conductivity of the swamp was estimated to be 1.5×10⁻² cm/s. This value is two to three orders of magnitude larger than that measured in a laboratory using small scale sediment core samples collected in the swamp. These results suggest that the presence of crowded, intricate and large animal burrows as well as sediment layers rich in mangrove humus increases permeability in the mangrove swamp. Further, it is suggested that the mangrove topography with the steep bank of the tidal creek plays an important role which enhances material exchanges through groundwater between the mangrove swamp and the adjacent offshore waters.     

The mechanism of inhibition of Ran-dependent nuclear transport by cellular ATP depletion

Rran-dependent nuclear transport requires a nuclear pool of RanGTP both for the assembly of export complexes and the disassembly of import complexes. Accordingly, in order for these processes to proceed, Ran-dependent nuclear import and export assays in vitro require the addition of GTP to produce RanGTP. Notably, no ATP requirement can be detected for these transport processes in vitro. But in vivo, when cells are depleted of ATP by the addition of sodium azide and 2-deoxyglucose to block ATP production by oxidative phosphorylation and glycolysis, respectively, Ran-dependent nuclear import and export are rapidly inhibited. This raised the question of whether there is an ATP requirement for these nuclear transport pathways in an intact cell that has remained undetected in vitro. Here we report that the free (but not total) GTP concentration rapidly drops to an undetectable level upon ATP depletion as does the availability of RanGTP. Our conclusion is that the inhibition of Ran-dependent nuclear transport observed upon ATP depletion in vivo results from a shortage of RanGTP rather than the inhibition of some ATP-dependent process.     

黄河三角洲潮沟形态特征对湿地植物群落演替的影响

通过野外调查明确黄河三角洲潮上带、潮间带、潮下带内潮沟的形态特征及植物群落演替特征, 并在此基础上探讨潮沟对植物群落的影响。结果表明: 1)潮沟共包括4级, 沿海到内陆方向, 呈现出潮沟分叉增加、级别增多、曲率变大的发育趋势。潮上带和潮下带均出现较大的宽深比, 潮间带的潮沟密度和分汊率较大; 2)共统计到植物10科15属15种, 大部分为草本植物。植物演替的方向为碱蓬-柽柳-芦苇和香蒲, 物种生态位宽度最大值随着演替的进行, 由碱蓬变为柽柳再到芦苇和香蒲; 3)潮沟曲率、潮沟密度和潮沟宽度与群落演替方向呈正相关, 而潮沟分汊率和潮沟深度与群落演替方向呈负相关。植物群落演替对潮沟分汊率的变化的反应最敏感, 潮沟密度次之。