Monitoring tagged plants, periods of maximum growth in length of 3.6 to 3.4 cm month1 were reported for April and July 1973, respectively in the Great Bay Estuary System of New Hampshire-Maine (Mathieson et al. 1976). Distinct yearly variations were found both in time of/maximal growth and rate of growth. Maximum growth rates were 2.3 and 2.2 cm month-1 in May and October respectively of the second measurement year (opp. cit.). An average of 0.8 cm month-1 was recorded for a Maine population by Vadas & Keser (1972). Three successive years of growth (elongation) measurements of Ascophyllum nodosum in the upper Ascophyllum zone at a site in Maine were 17.4+4.7, 11.5+5.5 and 12.2+5.5 cm (Keser and Larsen, 1984). Breton Provencher (1976) observing A. nodosum elongation in the St. Lawrence river estuary reported 26.4 cm and 24.8 cm in successive years. Nor-mandeau Assoc. Inc. (1977) found spring maximum growth rates of 2.2 cm month-1 and winter minimums of 1.2 cm month-1 in Maine. MacFarlane (1932) measured inter-vesicle length of Nova Scotian populations and calculated growth rates of 0.33 to 1.2 cm/month.Maximum rates of shoot elongation were recorded in the central part of the Ascophyllum zone but differences between zones did not exceed annual variation (Keser and Larsen, 1984). The length of the most distal internode (growth in year from formation of last vesicle) on unbroken primary shoots reaches a maximum at sites of intermediate wave exposure (Cousens, 1982). Growth rate increases with the length of intact primary shoots up to 15 cm at more exposed sites and 70 cm at more sheltered sites (Cousens, 1981a). The extremes of exposure were suggested to be suboptimal for growth due to conditions of desiccation/water turnover, and light (opp. cit.).A relative growth rate comparison was made between Ascophyllum nodosum and Fucus vesiculosus over seven-day periods at depths of 0 to 4 m. Growth in length and fresh weight decreased with depth except except for an increase near compensation depth (Ramus et al. 1977). Ascophyllum nodosum and F. vesiculosus grew in the same pattern but A. nodosum grew at a higher rate (opp. cit.). Maximum photosynthetic activity occurs in tissues of 2-3 years old in shoots with an average age of five to nine years (Khailov, 1976a). This peak of photosynthetic activity was correlated to the maximum dry weight for the 2-3 year age class of tissues (opp. cit.).A series of unique growth studies was conducted using short term measurements of apical growth in Ascophyllum nodosum with laser diffraction. Irradiance and temperature were correlated with hourly measurements of the growing tip to 1 m m. At artificial light of 12.5 Wm-2 and below, plants showed a direct relationship between growth and irradiance (Stromgren, 1977). Irradiance of 35 to 40 Wm-2 resulted in a reduced growth rate after 21 day periods. The fucoid species compensation point of 3 Wm-2 is only reached in the most extreme low light conditions during winter overcast days under a canopy (Schonbeck and Norton 1980). The saturation point 30-50 Wm-2 is not reached at the surface under similar conditions (opp. cit.). Response of apices to temperature was rapid between 2.5 to 30C for the first few hours, however, experiments of two to three weeks duration showed the optimal temperature or growth was below 17C (Stromgren, 1976). These laboratory studies correspond to field observations of maximum growth rates in moderate (6-10C) temperatures and light 150-250 1y d-1 (Mathieson et al. 1976). Elongation of Ascophyllum apices is linearly correlated with rapid air temperature increases to 35C over 1 to 4 hours (Stromgren, 1983). The duration of exposure at low tide and ambient air temperature can account for a large portion of shoot elongation and annual variability in growth.Growth continues during low nutrient conditions because of direct use of nitrogen from seawater during low ambient nitrogen levels and optimal light (Asare and Harlin, 1983). Tank cultured Ascophyllum nodosum growth is in agreement with previous lab and field observations reaching a maximum of 2.5% d-1 (length) at 15C. Growth was a linear function of light up to 20-30 yEm-2 5-1 (Fortes and Luning, 1980).Growth of zygotes settled on stone and brick substrata and transferred to the sea after ten days was found to be only 1 to 0.2 cm after one year and 0.5 to 1.5 cm after the second year (Sundene; 1973). Maximum growth rates for young Ascophyllum nodosum (10-80 mm) were 7 mm, 5 mm and 10 mm for the periods Sept.- Nov., Nov.-Feb. and Feb.-May in Scotland (Schonbeck and Norton, 1980). Studies of thallus growth immediately after germination in culture showed an increase with light intensity to 10,000 lux with temperature to 20C (Sheader and Moss, 1975). After 30 days, the thallus was 0.27 mm, corresponding to a growth of 0.32 cm year-1 similar to the in-situ values of Sundene, 1973.4.2 ProductivityIn a gradient of wave exposure, annual production (measured by standing crop differences) increases with wave shelter (Cousens, 1981b), Table 1. However the maximum production per unit biomass occurred on semi-exposed locations corresponding with an increase of reproductive effort. The turnover time for vegetative biomass ranged from 2.9 years at semi-exposed sites to 10.98 years at extreme wave exposed sites. Annual production varied by a factor of 2 within a 10 km section of coast line (opp. cit.).Table 1. Standing crop and production (kg dry wt m-2 at sites of different exposures to wave action. Sites 1-6 Polly Cove, Site 7 Dover Soi. P/B was calculated from the Modified Baardseth production estimates. (Cousens 1981b).Site1234567April Standing0.671.501.392.043.253.953.52Crop (+2 s.e.)+0.34+0.07+0.09+0.13+0.25+0.97+0.61Modified Baardseth method0.150.351.101.251.371.661.72Standing Crop0.050.260.950.940.841.071.05Difference Method P/B0.220.710.790.610.420.420.49Differences between the annual maximum and minimum standing crops obtained in seasonal surveys of Ascophyllum nodosum biomass were used to calculate annual production for a number of widely separated sites (Cousens 1984, Table 2). The maximum value of annual production was found in southwestern Nova Scotia equivalent to 1,015 g c m-2 (Cousens, 1981a).Table 2. Annual production and production to biomass ratios for Ascophyllum nodosum populations (Cousens 1964).Locationt dry wt ha-1P/BNew England15.86White Sea13.65Spain23.64Nova Scotia (pre-harvest)20-26.33-.25Nova Scotia6.1-28.2*.22-.79*Corrected for growth loss prior to sampling Cousens 1964.4.3 BiomassBiomass density in southwestern Nova Scotia is among the highest in the world (Table 3). Maine and southwestern Nova Scotia values are similar but Norwegian biomass density is closer to the levels in the St. Lawrence Estuary (Table 3). Survey methodology varies greatly between studies from selected samples in the Ascophyllum zone (McFarlane, 1952; Topinka et al. 1981) to a volume "eyeball" estimate (Pielou, 1981).Table 3. Biomass density of Ascophyllum nodosum populations in the North Atlantic. Source 

 Location Tonnes/Hectare wet weight McFarlane 1952 Southwestern Nova Scotia 198a Sharp 1981 Southwestern Nova Scotia 120  110b Cousens 1981a St. Margarets Bay Nova Scotia 96  45a Breton- Provencher 1976 St. Lawrence Estuary Quebec 20  5a Topinka et al 1981 Lincoln County Maine 80ac Chock & Mathieson 1976 Great Bay Estuary New Hampshire 72a Keser et al 1981 Average of six sites Boothbay, Maine 84  60a Baardseth 1970 South Norway 6.9  1.3 

 Central Norway 24.1  4.0 

 North Norway 14.4  1.7 Munda 1978 Iceland 69 aunharvested stock 

bharvested stocks 

cincludes all FucoidsSurveys and spot biomass values on the Atlantic coast of Nova Scotia and New Brunswick were combined with existing air photo analysis to estimate standing crops for Ascophyllum nodosum (Smith and Loucks, 1980). Some areas, e.g. the Nova Scotia Eastern Shore, have no ground truthing and other areas had poor quality air photos due to high tides. Taking a conservative estimate from the range of probable values the Atlantic and Fundy coasts of Canada support 442,000 t of A. nodosum wet weight (Table 4). A standing crop of 180,000 tons for southwestern Nova Scotia was estimated prior to harvesting and was based on selective sampling methods (McFarlane, 1952).Mean fucoid biomass per meter of a shoreline in Maine was 352.17 kg m-1 versus 540.0 kg m-1 in southwestern Nova Scotia (Topinka et al., 1981). The primary difference between these areas was a bed width of 45 m in Nova Scotia versus 22 m in Maine.Seasonal variation in Ascophyllum biomass was significant at a New Hampshire site (Chock and Mathieson, 1983). Peak biomass occurred in August 105-135 g 0.1m-2 and declined to 25-42 g 0.1m-2 in September. However, at a southern Nova Scotian site the only significant change in standing crop occurred (Fig. 8) in June-July after the loss of receptacles (Cousens, 1981a).Table 4. Total standing crop of A. nodosum from the Atlantic and Bay of Fundy shores of Nova Scotia and New Brunswick (Smith and Loucks 1980).Marine Plant District AreaStanding crop wet tFucoids estimated harvestable annual yield wet t14, Southern New Brunswick51,0003,80012, Minas Basin to Chebogue42.000+4,20012, Chebogue to Cape Sable Island125.000+15,00012, Cape Sable100.000+10,00011, Medway to Chedabucto Bay154,00011,500Total Atlantic & Fundy Coasts of Mainland Nova Scotia & New Brunswick472,00044,500From Smith & Loch 1980 

 + - Discounted by the author due to sampling methodology and assumptions of estimate.Figure 8. Seasonal variation in standing crop (dry weight) at Polly Cove, southeastern Nova Scotia. The same plants were measured each time by a non-destructive method (Pielou, 1981). Vertical line indicates the mean  2 s.e. (Cousens, 1981a).Biomass density decreases at both extremes of its vertical distribution on the shore (Chock and Mathieson, 1983; Cousens 1981a; Pringle and Semple, 1980) (Fig. 4). The spatial distribution of biomass is related to substratal types. In an area of mixed rock and gravel, patches of Ascophyllum averaged 1.5 to 2 m in maximum dimension. Rock and cobble substrata supported larger 2.0-2.5 m patches of A. nodosum (Black1, unpublished data). Biomass density increases with greater quantities of stable substrata (Fig. 9) (Topinka, 1980).R.Black. Univ. Western Australia, Perth, Australia.5. HARVESTING, BIOMASS RECOVERY AND ECONOMICS 5.1 Harvesting

 5.2 Regrowth

 5.3 Economics of harvesting techniques

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