Behaviour of adult Pacific lamprey in near‐field flow and fishway design experiments

Abstract An experimental fishway was used to examine adult Pacific lamprey, Lampetra tridentata (Gairdner), behaviour in a series of attraction and passage‐performance tests. Among all experiments, lamprey oriented to the fishway floor and walls and were attracted to both ambient and concentrated flow. When confronted with high‐velocity areas (vertical‐slot and submerged‐orifice weirs), many lamprey failed to pass upstream. However, lamprey were able to find and take advantage of low‐velocity refuges when they were provided. Lamprey climbed shallow‐ and steep‐angled ramps when attraction cues were sufficient and other passage routes were restricted. The combined results demonstrated the passage challenges that fishways designed and operated for salmonids present to non‐salmonid species. They also highlight the importance of evaluating trade‐offs between fishway attraction and passage efficiency. The experiments were integrated with tagging studies and development of lamprey‐specific passage structures, a research combination that provided an effective template for fishway performance evaluations.     

Climbing success of adult Pacific lamprey on a vertical wetted wall

The ability of adult Pacific lamprey, Entosphenus tridentatus (Gairdner), to climb has enabled development of lamprey‐specific fishways. Sheer vertical climbing elements are needed for these structures when fishway installation space is limited. A 1.6‐m high aluminium wetted wall was built to test Pacific lamprey vertical climbing in relation to flow and three water delivery mechanisms: overflow, upwelling and sidewelling. Passage success and passage times were recorded for individual Pacific lamprey (n = 122). For those that interacted with the structure (n = 104), 94% successfully ascended the wall (76% on their first attempt). Average time from first interaction with the structure to exit ranged from 19.5 to 47.0 min. Success rates and passage times were not significantly affected by flow or water delivery treatments. This indicates that vertical elements can facilitate lamprey passage in the 0.13–3.59 L min^?1 cm^?1 flow range tested and may greatly expand the design alternatives in certain situations.     

Effects of water velocity,turbulence and obstacle length on the swimming capabilities of adult Pacific lamprey

The swimming capabilities of adult Pacific lamprey, Entosphenus tridentatus Gairdner, were tested in an experimental vertical‐slot fishway in response to three different fishway features: water velocity (1.2, 1.8 and 2.4 m s^?1), the length of the vertical slot (0.33, 0.66, 1.00 m) and during the presence/absence of a turbulence‐intensifying structure that increased turbulence for each respective treatment. Passage success was lowest (52.7%) and attachment rates were highest (92.9%) during the most difficult passage conditions (high velocity, high turbulence, long slot length). Passage success was >83% for all other treatment combinations. Although passage of the vertical‐slot weir was not associated with fish body size, Pacific lamprey with larger dorsal distances (distance between the two dorsal fins) were more likely to pass the vertical‐slot weir. Increased attachment rates and longer attachment times during strenuous passage conditions suggest that endurance capacity may be an important factor limiting the passage of Pacific lamprey at fishway obstacles.     

Factors affecting dam passage and upstream distribution of adult Pacific lamprey in the interior Columbia River basin

Low dam passage rates of adult Pacific lamprey (Entosphenus tridentatus) during upstream spawning migration have been implicated in the species' steep decline in the interior Columbia River basin. In this 2000–2010 study, we used radiotelemetry to evaluate potential predictors of lamprey passage success at McNary Dam, located ~469 river kilometres (rkm) from the Pacific Ocean. The tagged population included 276 lampreys collected at McNary Dam and 53 collected at Bonneville Dam (rkm 235) that migrated to McNary Dam. McNary Dam passage efficiency was similar for the two samples, with multiyear estimates of 0.65 and 0.75, respectively. Larger‐bodied lampreys and those with earlier migration timing were more likely to return to McNary Dam after release, to pass the dam and to be detected upstream from McNary reservoir. Far more lampreys entered the upper Columbia River than the Snake River, suggesting that environmental cues (e.g., water discharge, temperature) or conspecific cues (e.g., pheromone concentrations) affect lamprey distribution above this large confluence. Overall, results indicate that Pacific lamprey passage success at barriers depends on a combination of individual lamprey traits plus seasonal and site‐specific effects on behaviour.     

Role of large‐ and fine‐scale variables in predicting catch rates of larval Pacific lamprey in the Willamette Basin,Oregon

Pacific lamprey Entosphenus tridentatus is an anadromous fish native to the Pacific Northwest of the USA. That has declined substantially over the last 40 years. Effective conservation of this species will require an understanding of the habitat requirements for each life history stage. Because its life cycle contains extended freshwater rearing (3–8 years), the larval stage may be a critical factor limiting abundance of Pacific lamprey. The objective of our study was to estimate the influence of barriers and habitat characteristics on the catch‐per‐unit‐effort (CPUE) of larval Pacific lamprey in the Willamette River Basin, Oregon, USA. We sampled lampreys at multiple locations in wadeable streams throughout the basin in 2011–13 and used an information theoretic approach to examine the relative influence of fine‐ and large‐scale predictors of CPUE. Pacific lamprey was observed across the basin, but its relative abundance appeared to be limited by the presence of natural and artificial barriers in some sub‐basins. Lower velocity habitats such as off‐channel areas and pools contained higher densities of larval lamprey; mean Pacific lamprey CPUE in off‐channel habitats was 4 and 32 times greater than in pools and riffles respectively. Restoration and conservation strategies that improve fish passage, enhance natural hydrologic and depositional processes and increase habitat heterogeneity will likely benefit larval Pacific lamprey.     

Using sprint swimming performance to predict upstream passage of the endangered Macquarie perch in a highly regulated river

Abstract Barriers to migration are a major threat to freshwater fish populations. A novel measurement of swimming speed performance and an analytical model were used to calculate the probability that endangered Macquarie perch, Macquaria australasica Cuvier, can swim through instream structures. Over the range of river discharges (2.7–166 ML day^?1) and temperatures (10–22 °C) examined, M. australasica could successfully pass through a rock‐ramp fishway and nearby natural riffles under most river discharges, whilst pipe culverts were passable only to large individuals (>17.7 cm total length) under a high river discharge. Cold water temperatures (below 16 °C) significantly reduced the likelihood of passage in each case. It was concluded that both the volume and thermal characteristics of environmental flow releases should be considered when assessing and remediating potential instream barriers to fish passage in regulated river systems.     

Modelling the impacts of environmental variation on habitat suitability for Pacific saury in the Northwestern Pacific Ocean

We developed habitat suitability index (HSI) models for two size classes of Pacific saury Cololabis saira in the Northwestern Pacific Ocean. Environmental data, including sea surface temperature, sea surface height, salinity, and net primary production, and catch and effort data from Taiwanese distant‐water stick‐held dip net fisheries during the main fishing season (August–October) during 2002–2015 were used. Habitat preferences and suitable habitat area differed between size classes. The suitable habitat was located between 40–47.5°N and 145–165°E for large‐sized Pacific saury but encompassed a greater area (35–47°N and 140–165°E) for medium‐sized Pacific saury. Both size classes were affected by substantial interannual variation in the environmental variables, which in turn can be important in determining the potential fishing grounds. We found a significant negative relationship between the suitable habitat area and the Niño3.4 indices with a time‐lag of 6 months for the large‐sized (r = ?0.68) and medium‐sized (r = ?0.42) Pacific saury, respectively, as well as the total landings of Pacific saury by all fishing fleets (r = ?0.46). As remotely‐sensed environmental data become increasingly available, HSI models may prove useful for evaluation of possible changes in habitat suitability resulting from climate change or other environmental phenomena and in formulating scientific advice for management.     

The feeding ecology of larval Pacific hake (Merluccius productus) in the California Current region: an updated approach using a combined OPC/MOCNESS to estimate prey biovolume

A combined optical particle counter (OPC) and multiple opening and closing net and environmental sensing system (MOCNESS) was used to obtain simultaneous measurements of the fine‐scale distribution of ‘prey‐sized’ particles and the vertical distribution of larvae of Pacific hake (Merluccius productus). Physical properties were also measured. The data were used to describe the feeding ecology of Pacific hake larvae, and to explore the effects of prey abundance, time of day, temperature and depth on feeding. Pacific hake larvae consumed a wide variety of prey including copepod eggs, nauplii, copepodites, and euphausiid metanauplii. Calanoid copepodites comprised > 75% of the ingested prey volume. First‐feeding larvae were 2.5–3.0 mm SL. These larvae consumed prey 40–200 μm wide. Larvae 3.0–6.5 mm long ingested prey 40–400 μm wide and larvae > 6.5 mm long ingested prey 400–700 μm wide. There were clear diel patterns in feeding incidence and prey volume ingested. Feeding commenced between 06:01 and 08:00 hours PST and continued until ～ 16:00 hours. Ingested prey items remained in the gut until complete gut evacuation occurred near dawn. The volume of food ingested was estimated using two indices, the cube root of the prey volume (CRPV) and the cube root of the stomach volume (CRSV). Standard length, log elapsed time since gut evacuation, depth and particle biovolume contributed significantly to variation in both indices. Temperature did not contribute to variability in either CRPV or CRSV.     

Forage and migration habitat of loggerhead (Caretta caretta) and olive ridley (Lepidochelys olivacea) sea turtles in the central North Pacific Ocean

Satellite telemetry from 26 loggerhead (Caretta caretta) and 10 olive ridley (Lepidochelys olivacea) sea turtles captured and released from pelagic longline fishing gear provided information on the turtles’ position and movement in the central North Pacific. These data together with environmental data from satellite remote sensing are used to describe the oceanic habitat used by these turtles. The results indicate that loggerheads travel westward, move seasonally north and south primarily through the region 28–40°N, and occupy sea surface temperatures (SST) of 15–25°C. Their dive depth distribution indicated that they spend 40% of their time at the surface and 90% of their time at depths <40 m. Loggerheads are found in association with fronts, eddies, and geostrophic currents. Specifically, the Transition Zone Chlorophyll Front (TZCF) and the southern edge of the Kuroshio Extension Current (KEC) appear to be important forage and migration habitats for loggerheads. In contrast, olive ridleys were found primarily south of loggerhead habitat in the region 8–31°N latitude, occupying warmer water with SSTs of 23–28°C. They have a deeper dive pattern than loggerheads, spending only 20% of their time at the surface and 60% shallower than 40 m. However, the three olive ridleys identified from genetics to be of western Pacific origin spent some time associated with major ocean currents, specifically the southern edge of the KEC, the North Equatorial Current (NEC), and the Equatorial Counter Current (ECC). These habitats were not used by any olive ridleys of eastern Pacific origin suggesting that olive ridleys from different populations may occupy different oceanic habitats.     

Sea lamprey avoid areas scented with conspecific tissue extract in Michigan streams

Three in‐stream experiments were conducted to determine whether sea lamprey, Petromyzon marinus L., tissue extract (alarm cue) and 2‐phenylethylamine hydrochloride (PEA HCl, a putative predator cue) influenced the distribution of migrating adult sea lamprey. Experiments evaluated sea lamprey movement when an odour was applied to (1) a tributary of a larger stream; and (2) half of a stream channel. Fewer sea lamprey entered the tributary and side of the river scented with sea lamprey tissue extract compared to the control treatment. Sea lamprey did not avoid the tributary and side of the river scented with PEA HCl. A final laboratory experiment found no difference in the avoidance response of sea lamprey to PEA HCl mixed with river water vs PEA HCl mixed with water from Lake Huron. As such, the lack of sea lamprey response to PEA HCl in the stream was unlikely to have been caused by the presence of the river water. Rather, the difference between laboratory and field results may be attributed to the complexity of the physical environment.     

Abiotic triggers for sea and river lamprey spawning migration and juvenile outmigration in the River Rhine,Germany

The activity patterns of migrating lampreys (sea lamprey Petromyzon marinus and river lamprey Lampetra fluviatilis) are poorly understood. This study uses extensive screening data from the cooling water intake of a nuclear power plant on the River Rhine, Germany, gathered on 1308 days over 46 consecutive months between 2011 and 2014. More than 34,000 lampreys were retrieved, of which the great majority (99.9%) were outmigrating nonfeeding postmetamorphic individuals. The main factors controlling outmigration were season, water temperature and river discharge. The bulk of downstream migration was observed each year in spring at water temperatures between 9 and 12°C, with earlier migrations during significant discharge peaks between December and February. The migration activity of postmetamorphic individuals declined during summer and autumn, regardless of discharge. Turbidity increased the occurrence of postmetamorphic lampreys and supported diurnal migration, which accounted for nearly 20% of all lamprey records. In winter, outmigrating individuals of both species exhibited significantly longer body lengths than those detected during other seasons. Haematophagus postmetamorphic sea lampreys were also retrieved (n = 28) and represent the most upstream records of parasitically feeding sea lamprey detected thus far. Adult lampreys migrating upstream were found mainly in spring, but in very low numbers, indicating high prespawning mortalities. The study points to a challenging future for the protection of lampreys in this area: the long duration and the round‐the‐clock nature of the observed migration demand flexible solutions to realise safe operational windows for water intake stations and to minimise impacts on juvenile lampreys.     

Development of diets for the intensive culture of Pacific lamprey

The Pacific lamprey Entosphenus tridentatus is an ancestral species of critical importance to the ecosystem and indigenous cultures in the Pacific Northwest. Conservation aquaculture has been proposed as a potential technique to restore Pacific lamprey populations. Intensive culture methods and diets for this species have not been developed. A sixteen week feeding trial tested the effects of seven diet treatments on the survival, growth, fatty acid profile and whole body lipid content of Pacific lamprey ammocoetes. Dietary treatments were: active dry yeast, yeast plus fish oil emulsion, micro‐algae, micro‐algae plus fish oil emulsion, yeast with micro‐algae, yeast with micro‐algae plus fish oil emulsion and yeast with larval fish diet. Each diet was offered to five replicate tanks stocked with 20 ammocoetes that were 51 days post hatch. Survival during the trial was not affected by diet. The greatest length and weight increases were in fish fed diets containing yeast. Growth decreased as the amount of algae in the diet was increased. Lipid retention was significantly higher in fish fed yeast with larval fish diet relative to the other treatments. Feed conversion ratio was lowest in fish fed diets containing yeast. Whole body fatty acid profiles tended to reflect the fatty acid profile of the diet. Percentages of 20:5n‐3 and 22:6n‐3 were significantly higher in fish fed diets containing fish oil emulsion. Overall, yeast with larval fish diet provided the best growth performance in larval Pacific lamprey.     

Marine biogeochemical response to a rapid warming in the main stream of the Kuroshio in the western North Pacific

Impact of climate change on marine biogeochemical parameters and ecosystem is one of the important issues of our environment. Direct evidence of marine pelagic ecosystem changes is found with warming of sea water and sea‐level rise in the main stream of the Kuroshio in the East China Sea and the western North Pacific during these three decades based on the analysis of long‐term comprehensive hydrographic observations. In terms of annual mean, the warming rate of surface air temperature and sea surface temperature ranged from 0.15 to 0.21°C per decade in and around the main stream of the Kuroshio in the East China Sea, which exceed the global mean warming rate of 0.128 ± 0.026°C per decade during the period from 1956 to 2005 reported in IPCC 2007. One of the features in this rapid warming region is an increase of number of Pterosagitta draco, a cosmopolitan warm‐water zooplankton. Biogeochemical parameters, such as wet weight of zooplankton, plant pigment and nutrients concentration in the upper 200 m have been decreasing while dissolved oxygen content and seawater temperature have been increasing in the upper 200 m in the main stream of the Kuroshio in the East China Sea. These observed linear trends of the biogeochemical parameters would be foresights for temperate oceans in the future.     

From shallow to deep waters: habitats used by larval lampreys (genus Petromyzon and Lampetra) over a western European basin

Abstract –  Habitat requirements of lamprey ammocoetes (Petromyzon marinus and Lampetra genus) were investigated, for the first time, from shallow to deep waters, at different spatial scales across the Gironde‐Dordogne continuum, thanks to a water suction dredge. Fish‐habitat relationships were assessed through two complementary statistical analyses: habitat‐use curves and habitat suitability models using the Boosted Regression Trees (BRT) technique. Analyses were performed on a small‐size data set that was characterised by the low prevalence of lamprey. The sea lamprey larvae occurred in deeper areas than their Lampetra genus counterparts. ‘Pools’ of 2 m’ depth and more were optimal habitats for the former species. Among the environmental variables retained to model lamprey occurrences, the mesohabitat (a categorical variable) was demonstrated to be highly influential, in terms of fine grain‐size substratum and vegetation cover. These preliminary results suggest that monitoring using the water suction dredge method may contribute to sea lamprey conservation.     

Gauging weirs impede the upstream migration of adult river lamprey Lampetra fluviatilis

Abstract The ability of individual and groups of 30 migrating adult river lamprey Lampetra fluviatilis L. to pass a Crump or flat‐v gauging weir under two discharge regimes (moderate and low) was assessed in an experimental channel. Despite repeated attempts by the lamprey, the Crump weir remained impassable during all trials. Lamprey passage over the flat‐v weir occurred only during group trials at low discharge (5.73 ± 0.19 L s^?1) and only as a single burst swimming event via the deeper water passing over the centre of the weir. Lampreys generally approached the weirs along the channel walls and particularly favoured the true right wall associated with elevated velocities during low discharge and reverse currents at moderate discharge (68.06 ± 2.41 L s^?1). Time spent immediately below the weir was lower than expected compared with further downstream. Rate of weir approach, attempts to pass (absolute number and as a proportion of the total approaches) and time spent immediately downstream of the weir were highest for the Crump weir at low discharge and the flat‐v weir at moderate discharge. The present study suggests that gauging weirs may severely impede the movements of migrating adult river lamprey under low to moderate discharges.     

Condition assessment of lamprey populations in the Yorkshire Ouse catchment,north‐east England,and the potential influence of physical migration barriers

• 1. River lamprey (Lampetra fluviatilis) and sea lamprey (Petromyzon marinus) are designated features of the River Derwent Special Area of Conservation (SAC) and the Humber Estuary (a possible SAC). This study determined the condition of lamprey populations in the Yorkshire Ouse catchment by assessing the species composition, distribution, abundance and size‐structure of larval (ammocoete) populations in five major tributary rivers (Derwent, Swale, Ure, Nidd and Wharfe).
• 2. According to EU condition assessment criteria, Lampetra populations (assumed to be mostly river lamprey) are at present in favourable condition, with site mean (±SE) densities ranging from 2.7 (±1.2) to 160.3 (±50.5) individuals m^?2 (all river means >2 individuals m^?2), and at least two size (≈age) classes present in optimal microhabitats. By contrast, no sea lamprey larvae were recorded, suggesting that populations of this species are in unfavourable condition.
• 3. Actions to protect and enhance nationally or internationally important stocks must be implemented from at least a catchment perspective, because many of the issues affecting such species are not localized. With respect to lampreys, particular attention should be given to protecting spawning and nursery habitats, improving water quality, reducing impingement at abstraction points, preventing exploitation at spawning grounds and increasing passage at potential physical obstructions.

Copyright © 2007 John Wiley & Sons, Ltd.     

Cumulative impacts of habitat fragmentation and the environmental factors affecting upstream migration in the threatened sea lamprey,Petromyzon marinus

1. River ecosystems are often fragmented by artificial structures, such as weirs. For anadromous species, these structures can impede access to upstream spawning sites and ultimately lead to severe population declines.
2. This study focused on the freshwater spawning migration of the sea lamprey, Petromyzon marinus, an anadromous species threatened by habitat fragmentation across its native range. To quantify the cumulative impacts of multiple weirs on upstream-migrating adults, and to explore the environmental factors affecting migratory movements, passive acoustic telemetry was applied to 56 individuals during their spawning migration in the heavily fragmented River Severn basin, UK.
3. While 89% of tagged sea lamprey passed the first weir upstream of the release site on the main river, only 4% passed the fifth weir. For 85% of migrants, the upstream extent of migration was immediately downstream of a weir. Individuals that passed weirs upstream of the release site (n = 50) took 21.6 ± 2.8 days to reach their most upstream location, experiencing cumulative passage times at weirs of 15.7 ± 2.8 days; these delays constituted a median of 84% of total upstream movement times.
4. Multistate models showed that the weir passage rates of sea lamprey in tidal and non-tidal areas increased significantly when downstream river level and discharge were elevated. Upstream-to-downstream changes in direction were frequent downstream of weirs, but rare in unobstructed river sections.
5. The results provided evidence for a cumulative effect of multiple weirs on sea lamprey movements, substantially delaying upstream migrants and limiting their spawning to atypical habitat. The results also demonstrated the crucial roles of high tides and elevated discharge events in enabling weir passage. Although the Severn Estuary features conservation designations for sea lamprey, this study reveals that barriers are inhibiting their upstream migration, a problem that should be addressed to assist sea lamprey conservation.

     

What goes up must come down: Diel vertical migration in the deep‐water sablefish (Anoplopoma fimbria) revealed by pop‐up satellite archival tags

The sablefish (Anoplopoma fimbria) is a long‐lived species with wide distribution throughout the North Pacific Ocean. While adult sablefish are considered a deep‐water fish, diet analyses suggest that they undergo vertical migrations that could be related to prey movement and feeding. Pop‐up satellite archival tags (PSATs) were used to observe the fine‐scale depth selection behavior of adult sablefish tagged off the Washington coast during the summer from June to August. Tags were physically retrieved after they surfaced using direction‐finding equipment so that complete datasets over the entire deployment were obtained from 14 tags. PSATs that recorded depth and temperature every 4 min during the deployment confirm that sablefish inhabit depths of 750 m or greater. However, a majority of the tagged fish underwent extensive vertical migrations that averaged 254.4 m overall and occurred at a 24 hr periodicity. Variations were observed among individuals in the amount of the deployment during which vertical migrations occurred, ranging from 12.37% to 63.48% of the time. During the vertical migration, fish ascended towards the surface at night and descended prior to daylight (i.e., diel vertical migration). Sablefish generally inhabited temperatures of 5°C but during the vertical migrations were found at temperatures from 6 to 10°C. Sablefish are opportunistic feeders with a large proportion of their diet being fish, euphausiids and cephalopods. Because these prey items also exhibit diel vertical migrations, it is possible that the vertical migratory behavior displayed by the sablefish was in response to the movements or the location of their prey.     

Northerly shift of warm‐water copepods in the western subarctic North Pacific: Continuous Plankton Recorder samples (2001–2013)

Spatial and temporal variation in copepod community structure, abundance, distribution and biodiversity were examined in the western subarctic North Pacific (40–53°N, 144–173°E) during 2001–2013. Continuous Plankton Recorder (CPR) observational data during the summer season (June and July) were analyzed. The latitudinal distribution of warm‐water species in June shifted northward after 2011 while no apparent latitudinal shift of cold‐water and other species was observed. Species number and the Shannon–Wiener biodiversity index (H′) in June tended to increase in the northern area after 2011. The warm‐water species abundance and center latitude of warm‐water distribution were positively correlated with sea surface temperature (SST) across sampling locations, whereas no significant correlations with SST were observed for cold‐water species or other species. Warm SSTs in June after 2011 appeared to cause the northward shift of warm‐water species distribution, which in turn contributed to the higher biodiversity in the northern area. This study demonstrated the rapid response of warm‐water species to warm SST variation, whereas cold‐water and other species did not exhibit such clear responses. These findings indicate that the response of copepods to environmental changes differs among copepod species, highlighting the importance of investigating lower trophic levels to the species level to evaluate individual species’ responses to climate change.     

Variability of the Pacific North Equatorial Current and its implications on Japanese eel (Anguilla japonica) larval migration

The zonal velocity produced by a Regional Ocean Modeling System (ROMS)‐based Pacific Ocean circulation model was validated against in situ measurements along the 137°E longitude. The Pacific model successfully reproduced the position and the shape of the North Equatorial Current (NEC) as well as the latitude of maximum surface velocity in the NEC region (8–17°N). The flow field produced by the Pacific model was then used to conduct numerical Lagrangian experiments, in which passive particles were released along a transect (142.5°E, 12.5–17°N) that traverses the known Japanese eel spawning area, and the effects of NEC strength and bifurcation latitude on the particle advection in the northwest Pacific were studied. Our results suggest that, in the 20‐yr period (1993–2012), the variability of the currents alone can cause interannual variability of one order of magnitude in the Kuroshio Entrance (KE), the percentage of particles entering the Kuroshio, the range of which varies from 43% in 1997 to 6% in 2012. The yearly‐averaged KE is not sensitive to the NEC bifurcation latitude. Instead, it is controlled by the average zonal velocity of a fixed domain (125–143°E, 13.5–17°N) and related to a recently‐developed climate index, the Philippines–Taiwan Oscillation (PTO). During the positive phase of the PTO, the zonal velocity in the domain, hence the yearly‐averaged KE, increases, and the opposite is true in the negative phase of the PTO. Considering only the trajectories, diel vertical migrations (DVM) in the top 400 m do not significantly affect Japanese eel larval transport, as incorporating DVM schemes does not increase the KE.