Combined effects of seawater temperature and nutrient condition on growth and survival of juvenile sporophytes of the kelp Undaria pinnatifida (Laminariales; Phaeophyta) cultivated in northern Honshu, Japan

Recently, withering of farmed juvenile sporophytes of the kelp Undaria pinnatifida (Harvey) Suringar has led to reduced production of this species in northern Japan, possibly because of the high water temperature and low nutrient concentration in cultivation areas. This problem may be solved by introducing parental plants with greater tolerance to high temperature and low nutrient conditions. We examined the combined effects of various temperatures (15, 20, 24, and 27 °C) and nutrient availabilities (seawater enriched with 25 % PESI medium and non-enriched seawater) on the growth and survival of cultivated juvenile sporophytes (1–2 cm) collected from Matsushima Bay, Miyagi Prefecture in northern Japan and Naruto, Tokushima Prefecture in southern Japan. The relative growth rates of juvenile sporophytes from both locations were significantly greater at 15 and 20 °C than at 24 and 27 °C. The juvenile sporophytes cultured in enriched medium had significantly higher relative growth rates than those cultivated in non-enriched seawater. Dead juveniles were observed in non-enriched seawater at all temperatures and the survival percentage decreased with increasing seawater temperatures. Compared to the juvenile sporophytes from Matsushima Bay, those from Naruto showed greater tolerance to high temperature even under the low nutrient condition. These results suggest that the withering of juvenile sporophytes is caused by the combined effects of low nutrient availability and high temperature. A potential solution to this problem is to introduce ecotypes with greater tolerance to high temperature and low nutrient conditions from a warmer region of Japan.     

Effects of experimental thinning on the growth and maturation of the brown alga Undaria pinnatifida (Laminariales; Phaeophyta) cultivated in Matsushima Bay,northern Japan

In northern Japan, massive production of high-quality specimens of the kelp Undaria pinnatifida prior to the traditional harvest season is strongly needed to meet commercial demand. To address this need, we tested the effects of controlled thinning by cutting small plants on the growth and maturation of sporophytes cultivated in Matsushima Bay, northern Japan. In early December 2009, the stocking densities of cultivated kelps were thinned to 10 and 5 plants per 4 cm thread section in two experimental groups. In contrast, no thinning was conducted during cultivation until March 2010 in the control group (about 16–20 plants per 4 cm thread section). Morphological features, photosynthesis, and carbon and nitrogen contents were compared among the three groups. Compared to kelps of control group, total length, stipe width, the length and width of sporophyll, and dry weights of blade and sporophyll showed significantly higher values in kelps of the two experimental groups. However, stipe length of kelps of control group was significantly greater than those of the two experimental groups. The photosynthetic rates and nitrogen contents of kelps of the two experimental groups were significantly greater than those of the control from January to March. No significant differences occurred in all these parameters between kelps of the two experimental groups. Based on these results, the growth and maturation of U. pinnatifida sporophytes can be promoted greatly by experimental thinning, and this will improve production of high-quality specimens of kelps and increase economic returns before the traditional harvest season.     

Temperature requirements for the growth of young sporophytes of Undaria pinnatifida and Undaria undarioides (Laminariales, Phaeophyceae)

The relative growth rate of young sporophytes of Undaria pinnatifida (Harvey) Suringar and Undaria undarioides (Yendo) Okamura was examined in order to understand the difference in distribution of these two species around the coast of Japan. The optimal temperature for growth of both species was similar at 20°C and the upper critical temperature for growth was also similar, at 27°C for U. pinnatifida and 26°C for U. undarioides. Therefore, the optimal and upper critical temperatures for growth of the young sporophytes are not the main factors determining the distribution of each species. Next, the lower critical temperatures for growth were examined. For the young sporophytes of U. pinnatifida, the lower limit was less than 5°C while for those of U. undarioides it was 15°C. Thus, the difference in the lower critical temperature for growth between the two species was approximately 10°C. During the period of young sporophyte growth in the field, the temperature at the mouth of Ise Bay, Japan, where U. pinnatifida occurs, ranges from 12.7°C in December to 13.1°C in April, with a minimum of 7.9°C in February. Our experiments indicate that young sporophytes are able to grow throughout this period. The temperature off Hamajima, Japan, where U. undarioides occurs, ranges from 19.1°C to 14.8°C during the same time period. Again, young sporophytes are able to growth throughout this period, although minimum winter temperatures are only just high enough for growth. These natural temperature ranges during the growth season of the sporophytes agree well with the experimentally determined temperature requirements for growth of each species. Therefore, the difference between the two species in the critical temperature required for growth of the young sporophytes, especially in the low temperature range, is one of the major factors determining the distribution pattern of each species.     

Morphological and physiological differences among cultivation lines of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in a common garden experiment using a tank culture system

Undaria pinnatifida is grown for food and industrial materials worldwide; therefore, advanced breeding is needed to meet quality and productivity requirements. In this study, we examined regional lines of U. pinnatifida from five cultivation sites in Japan with different environmental conditions: Oga (OGA, the northern Sea of Japan coast), Hirota Bay (HRT, the northeastern Pacific coast), Matsushima Bay (MAT, the northeastern Pacific coast), Naruto (the Seto Inland Sea coast) and Shimonoseki (SIM, the southern Sea of Japan coast). The sporophytes of these lines were cultured in a tank culture system under controlled environmental conditions, and their morphological characteristics, nutrient uptake kinetics (V _max, K _s and V _max/K _s ), and carbon, nitrogen and phosphorus contents were determined. Sporophytes from MAT grew faster, whereas those from SIM were smaller than those from the other sites. Although the blade thickness of sporophytes cultivated in the sea significantly differs among cultivation sites in the previous study, there was no significant difference in blade thickness among the regional lines cultivated in the tank. Sporophytes from OGA had the greatest V _max/K _s values and significantly greater nitrogen contents than the other lines. Therefore, the morphological characteristics of MAT and SIM sporophytes, and the nutrient uptake kinetics of OGA sporophytes may have a genetic origin. This indicates that these lines may represent useful resources for selective breeding, with MAT sporophytes providing faster growth and OGA sporophytes being well-adapted to low-nutrient conditions.     

Compensation of the brown alga Undaria pinnatifida (Laminariales; Phaeophyta) after thallus excision under cultivation in Matsushima Bay, northern Japan

In recent years, the crop yield of cultivated Undaria pinnatifida (Harvey) Suringar has not been able to meet commercial demand and so advances in cultivation technology are strongly needed to increase production. Interestingly, cultivation work has shown the yield of U. pinnatifida sporophytes may be increased significantly by thallus excision, which may cause compensatory growth of the remaining tissues. To test this hypothesis and clarify correlative mechanisms, we examined seasonal morphological characteristics, photosynthetic rates, nutrient uptake rates, and carbon and nitrogen contents of U. pinnatifida sporophytes after thallus excision at a point 30 cm from the meristem in late February and compared these parameters with control kelps grown without excision, which were cultivated together in Matsushima Bay, northern Japan. Compared to control kelps, the length and dry weight of blades increased significantly after excision, and the growth phase was prolonged for about 1 month. The photosynthetic rates, nutrient uptake rates, and carbon and nitrogen contents of excised kelps were significantly higher than those of the controls. After vegetative growth stopped, resources accumulating in thalli were translocated significantly to sporophylls for maturation, indicating maturation was not negatively affected by thallus excision. These results indicate U. pinnatifida exhibits a very strong compensatory ability in response to thallus excision and consequently, the yield could be increased due to an increase in harvest frequency.     

Growth of Ecklonia cava (Laminariales, Phaeophyta) sporophytes transplanted to a locality with different temperature conditions

Transplanting experiments were carried out to determine whether the small type sporophytes with short stipe of Ecklonia cava Kjellman (Laminariales, Phaeophyta) growing in a locality with warm temperatures, change into larger type with a long stipe when transplanted to a locality with cooler temperatures. Juvenile E. cava sporophytes, having a stipe shorter than 5 cm long were collected from Tei in Tosa Bay (southern Japan) (seawater temperature 15–29°C) and transplanted to Nabeta Bay (central Japan) (seawater temperature 13–25°C), where larger type E. cava sporophytes characterized by long stipe (ca 1 m) grow. They were attached to artificial reefs at the sea bottom (9 m depth) in Nabeta Bay to monitor their growth. For comparison, juvenile E. cava sporophytes of almost similar size growing in Nabeta Bay were also transplanted in the same way to the same experimental site. Observations of growth of sporophytes from Tei and Nabeta were carried out monthly for 2 years from November 1995 to October 1997. The transplanted Tei and Nabeta sporophytes showed an increase in stipe length and diameter from winter to spring, whereas almost no increase was observed during summer and autumn. At the end of the study period, the stipe of Nabeta sporophytes reached 25.6 cm in length and 17.0 mm in diameter, whereas that of Tei sporophytes reached 11.1 cm in length and 11.2 mm in diameter. The primary blade length was 16.0 cm in Nabeta sporophytes, whereas it was 5.5 cm in Tei sporophytes. Thus, Tei sporophytes still remained smaller than Nabeta sporophytes even under the same environmental conditions.     

Compensatory abilities depending on seasonal timing of thallus excision of the kelp Undaria pinnatifida cultivated in Matsushima Bay, northern Japan

Improved cultivation technology for the kelp Undaria pinnatifida is greatly needed to increase production to meet increasing commercial demand. A previous cultivation trial indicated that the crop yield of U. pinnatifida sporophytes could be increased greatly by thallus excision in late February due to compensatory growth of the remaining tissues. To develop this potential new cultivation technology, it is essential to identify the time period during which this kelp can compensate and its physiological responses to thallus excision. In this study, U. pinnatifida sporophytes were excised at about 30 cm above the meristem at the beginning of January, February, March, and April, respectively. Morphological features, photosynthesis, nutrient uptake, and carbon and nitrogen contents of excised kelps were measured and compared with these parameters in control kelps grown without excision. Both experimental and control kelps were farmed together in Matsushima Bay, northern Japan. The kelps excised in January and February showed significant increases in the lengths and dry weights of the blade, photosynthetic rates, nutrient uptake rates, and carbon and nitrogen contents compared with the control kelps, and the growth phase was prolonged for at least 1 month. No significant increases were found in dry weights and carbon and nitrogen contents of sporophylls until early April, which indicated that the maturation period was delayed. At the end of this experiment, the nitrogen contents of sporophyll tissues formed after excisions were significantly lower than those of tissues formed before excisions. In contrast, the kelps excised in March and April showed no significant increases in morphological and physiological parameters compared with control kelps. These results suggest that U. pinnatifida sporophytes exhibited great compensation when excisions were conducted during the growing phase in January and February but not in March and April when the maturation phase had started. The regulation of resource allocation to growth and maturation after thallus excisions in January and February likely results in prolongation of the growth phase and maturation phase in excised kelps.     

Effects of temperature and nutrients on microscopic stages of the bull kelp (Nereocystis luetkeana,Phaeophyceae)

Warming ocean temperatures have been linked to kelp forest declines worldwide, and elevated temperatures can act synergistically with other local stressors to exacerbate kelp loss. The bull kelp Nereocystis luetkeana is the primary canopy-forming kelp species in the Salish Sea, where it is declining in areas with elevated summer water temperatures and low nutrient concentrations. To determine the interactive effects of these two stressors on microscopic stages of N. luetkeana, we cultured gametophytes and microscopic sporophytes from seven different Salish Sea populations across seven different temperatures (10–22°C) and two nitrogen concentrations. The thermal tolerance of microscopic gametophytes and sporophytes was similar across populations, and high temperatures were more stressful than low nitrogen levels. Additional nitrogen did not improve gametophyte or sporophyte survival at high temperatures. Gametophyte densities were highest between 10 and 16°C and declined sharply at 18°C, and temperatures of 20 and 22°C were lethal. The window for successful sporophyte production was narrower, peaking at 10–14°C. Across all populations, the warmest temperature at which sporophytes were produced was 16 or 18°C, but sporophyte densities were 78% lower at 16°C and 95% lower at 18°C compared to cooler temperatures. In the field, bottom temperatures revealed that the thermal limits of gametophyte growth (18°C) and sporophyte production (16–18°C) were reached during the summer at multiple sites. Prolonged exposure of bull kelp gametophytes to temperatures of 16°C and above could limit reproduction, and therefore recruitment, of adult kelp sporophytes.     

Temperature requirements for the growth and maturation of the gametophytes of Undaria pinnatifida and U. undarioides (Laminariales, Phaeophyceae)

Gametophytes of two Undaria species, U. pinnatifida and U. undarioides (Laminariales, Phaeophyceae), were studied to determine their water temperature requirements in order to understand their different distributions in Mie Prefecture, Japan. The optimal temperature for growth was 20°C for gametophytes of both species, and the upper critical temperature for growth was also the same for both species at 28°C. Therefore, the optimal and critical temperatures for growth of the gametophytes are not the main factors determining distribution. The optimal temperature for maturation of U. pinnatifida was approximately 10–15°C, whereas it was closer to 20–21°C for U. undarioides, a difference between these species of at least 5°C. In autumn and early winter, the seawater temperature at the mouth of Ise Bay, where U. pinnatifida is distributed, ranges from 21.6°C (October) to 12.7°C (December), and off Hamajima, where U. undarioides is found, the range is from 22.7°C (October) to 19.1°C (December). The seawater temperatures from October to December, which is the maturation season for the gametophytes, agreed well with the optimal temperature requirements for maturation of the gametophytes of both species. Thus the difference in the maturation temperature range of the gametophytes is a major factor determining distribution of these Undaria species along the Japanese coast.     

Coalescence in wild organisms of the intertidal population of Lessonia berteroana in northern Chile: management and sustainability effects

Lessonia berteroana is the subject of 85 % of the total harvest of Chilean brown seaweeds, representing close to 10 % of total kelp biomass worldwide harvested for alginic acid extraction. Frequent incidence of coalescent holdfasts was detected in natural populations, and this process gives rise to fused sporophytes. This study presents the coalescence events in natural populations in northern Chile. During 2011, in natural intertidal populations, 435 target sporophytes of different sizes were observed weekly and 63.90 % (278) showed physical signs of coalescent disks. We were able to distinguish five fusion modes depending on the number and size of each of the participant sporophytes and their spatial distribution. There was a progressive decrease of density and an increase in the number of stipes of each sporophyte over time. Two processes were recognized: active fusion of juvenile sporophytes and passive fusion of adult and senescent sporophytes. However, most of the coalescence processes were detected in juvenile sporophytes with a holdfast diameter of 0.5 to 2 cm. The minimum distance between pairs of coalescing sporophytes was 0.5 cm, and the maximum distance was 13 cm. For good harvesting practices, it is recommended that plants with over 20 cm holdfast diameter are harvested from natural populations because over this size, all of the plants have reproduced. Coalescence events would produce plants with legal size requirement; however, these plant units would not be reproductive, affecting the sustainability of this important coastal resource.     

The effect of irradiance and temperature responses and the phenology of a native alga,Undaria pinnatifida (Laminariales), at the southern limit of its natural distribution in Japan

Phenology, irradiance, and temperature characteristics of an edible brown alga, Undaria pinnatifida (Laminariales), were examined from the southernmost natural population in Japan, both by culturing gametophytes and examining the photosynthetic activity of sporophytes using dissolved oxygen sensors and pulse amplitude-modulated chlorophyll fluorometer (IMAGING-PAM). Our surveys confirmed that sporophytes were present between winter and early summer, but absent by July. IMAGING-PAM experiments were used to measure maximum effective quantum yield (ΦII at 0 μmol photons m^?2 s^?1) for each of 14 temperatures (8–36 °C). Oxygen production was also determined over a coarser temperature gradient. Net photosynthesis and ΦII (at 0 μmol photons m^?2 s^?1) were observed to be temperature-dependent; the maximum ΦII was estimated to be 0.67, occurred at 21.2 °C, and was nearly identical to the optimal temperature of the net photosynthetic rate (21.7 °C). A net photosynthesis–irradiance (P–E) model revealed that saturation irradiance (E _k) was 119.5 μmol photons m^?1 s^?1, and the compensation irradiance (E _c) was 17.4 μmol photons m^?1 s^?1. Culture experiments on the gametophytes revealed that most individuals could not survive temperatures over 28 °C and that growth rates were severely inhibited. Based on our observations, temperatures greater than 20 °C are likely to influence photosynthetic activity and gametophyte survival, and therefore, it is possible that this species might become locally extinct if seawater temperatures in this region continue to rise.     

Morphometric study of Ecklonia cava (Laminariales, Phaeophyta) sporophytes in two localities with different temperature conditions

Sporophytes of Ecklonia cava Kjellman (Laminariales, Phaeophyta) were collected seasonally from within 3–4 replicate, 1‐m² quadrates, haphazardly placed in dense assemblages at 6–9 m depth in Tei, Tosa Bay (southern Japan; water temperature 15–29°C) from 1995 to 1996, and in Nabeta Bay, Shimoda (central Japan; water temperature 13–25°C) from 1996 to 1997. Growth rings were checked for all samples and mean values of each biometric parameter of sporophytes 1‐year‐old and over were compared. The plant length (stipe length + primary blade length) was always shorter in Tei sporophytes (24–52 cm) than Nabeta sporophytes (70–100 cm), the difference being mainly owing to the shorter stipe length in Tei sporophytes (7–14 cm) than in Nabeta sporophytes (54–83 cm). However, the primary blade length was sometimes longer in Tei sporophytes (12–38 cm) than in Nabeta sporophytes (14–21 cm). Stipe diameter, longest bladelet length and primary blade width were mostly less in Tei sporophytes (8.6–12.4 mm, 5.5–7.0 cm and 23.4–38.0 cm, respectively) than Nabeta sporophytes (16.4–20.2 mm, 9.2–12.0 cm and 43.0–52.6 cm, respectively). Nevertheless, the number of bladelets of sporophytes from Tei (15–28) and Nabeta (18–29) were within the same range. At Tei, 32–43% of the sporophytes had wrinkled blades in summer and autumn, whereas wrinkled primary blades or bladelets were not observed in Nabeta sporophytes throughout the experimental period. These morphometric differences of E. cava between the two localities are suggested to be dependent on environmental factors, especially seawater temperature.     

Intraspecific Genetic Diversity of Undaria pinnatifida in Japan, based on the Mitochondrial cox3 Gene and the ITS1 of nrDNA

The intraspecific genetic diversity of the kelp Undaria pinnatifida (Harvey) Suringar (Laminariales, Phaeophyceae) was investigated using DNA sequences of the mitochondrial cytochrome oxidase subunit 3 (cox3) gene and internal transcribed spacer 1 (ITS1) of nuclear ribosomal DNA in plants collected from 21 localities along the Japanese coast between 2001 and 2003. Morphological variation was also examined and compared with the genetic diversity. Cox3 analyses of 106 plants revealed 9 haplotypes (I–IX) that differed from each other by 1–7 bp (all synonymous substitutions). Haplotype I was distributed in Hokkaido and the northern Pacific coast of Honshu, while haplotype III was found along the Sea of Japan coast of Honshu. Other types were found along the central and southern coast of Honshu. ITS1 analyses of 42 plants revealed 0–1.7% nucleotide differences, but plants from the Sea of Japan coast and northern Japan had similar sequences. The lower genetic differentiation along the Sea of Japan and northern coasts might be due to the recent establishment (after the middle of the last glacial period) of the Sea of Japan flora. The cox3 haplotype of cultivated plants was found in natural populations occurring close to cultivation sites (Naruto, Tokushima Pref., and Hokutan, Hyogo Pref.). This suggests that cultivated plants possibly escaped and spread or crossed with plants of natural populations. Morphological analyses of variation in 10 characters were conducted using 66 plants. The results showed no significant local variation owing to the wide variation in each population and did not support any forma previously described. No correlations between the morphological characters and cox3 haplotypes were detected.     

Geographic variation in cold hardiness of eggs and neonate larvae of the yellow-spotted longicorn beetle Psacothea hilaris

Cold hardiness of eggs and neonate larvae of the yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe) was examined using six geographical populations in Japan. Particular attention was paid to cold hardiness of eggs and neonate larvae of the subtropical population (Ishigaki), because the east Japan populations are considered to have been introduced from a subtropical area, and the overwintering stage in the east Japan populations is incidentally shifted from the original mature larval stage to the egg or neonate larval stages. When the eggs were exposed to low temperatures for 1 h, the decrease in hatchability became significant at –12°C in the southernmost two populations (Ishigaki and Naze), and at –16°C in the northern populations. After 1 h exposure to –20°C, few eggs could hatch in the Ishigaki population, whereas 27–55% of the eggs survived in the northern populations. Pre-chilling of the eggs at 10°C for 10 days enhanced the cold hardiness in all populations. This effect was particularly distinct in the subtropical population; the eggs of the Ishigaki population became as cold hardy as those of the northern populations after acclimation. These results suggest that the subtropical population is capable of establishing itself in east Japan, where the winter is cold.     

TEMPERATURE RESPONSES OF DISJUNCT TEMPERATE BROWN ALGAE INDICATE LONG-DISTANCE DISPERSAL OF MICROTHALLI ACROSS THE TROPICS1

We examined the temperature tolerance of microscopic phases from geographically disjunct isolates of eight species or closely related, putatively conspecific taxa of temperate brown algae with disjunct distributions. Maximum within-taxon differences were small and ranged from 1.6° to 4.3° C. Desmarestia aculeata and Sphaerotrichia divaricata, both with northern hemisphere amphioceanic distributions, showed little or no significant intraspecific variation between the mean upper survival limits (USL) of Atlantic and Pacific strains (δUSL ≤ 1.4°C), which would agree with a relatively recent separation of the respective populations. Among the plants with bipolar distributions, there was likewise very little difference (δUSL 0–1.1°C) between northern and southern hemisphere strains in Striaria attenuata and in the species pair Desmarestia viridis/D. willii. In Desmarestia ligulata, and in the species pairs Desmarestia firma/D. munda, Dictyosiphon foeniculaceus/D. hirsutus, and Scytothamnus australis/Scytothamnus sp., significant differences occurred, which indicate longer divergence times. δUSL in these cases ranged from 1.7° to 2.7°C, without overlap between strains from the northern and southern hemispheres. All species that passed the equator during cooler epochs had a USL of 26–27°C, at least in some geographical isolates. The NE Asian kelp Undaria pinnatifida, which passed the equator in recent times, had a USL of 29.6°C. We hypothesize that the mechanism of spreading in the amphipolar species studied was migration of vegetative microthalli. The more unlikely alternative hypothesis of continuous populations through the tropics during a cooler epoch would imply a drop in seawater temperatures to approximately 20° C in summer and 15° C in winter, which is not supported by paleoclimatic evidence.     

Thermal habitat of adult Atlantic salmon Salmo salar in a warming ocean

The year-round thermal habitat at sea for adult Atlantic salmon Salmo salar (n = 49) from northern Norway was investigated using archival tags over a 10 year study period. During their ocean feeding migration, the fish spent 90% of the time in waters with temperatures from 1.6–8.4°C. Daily mean temperatures ranged from −0.5 to 12.9°C, with daily temperature variation up to 9.6°C. Fish experienced the coldest water during winter (November–March) and the greatest thermal range during the first summer at sea (July–August). Trends in sea-surface temperatures influenced the thermal habitat of salmon during late summer and autumn (August–October), with fish experiencing warmer temperatures in warmer years. This pattern was absent during winter (November–March), when daily mean temperatures ranged from 3.4–5.0°C, in both colder and warmer years. The observations of a constant thermal habitat during winter in both warmer and colder years, may suggest that the ocean distribution of salmon is flexible and that individual migration routes could shift as a response to spatiotemporal alterations of favourable prey fields and ocean temperatures.     

Growth and survival rates of large-type sporophytes of Ecklonia cava transplanted to a growth environment with small-type sporophytes

Stipe lengths of sporophytes of Ecklonia cava Kjellman have been reported to be longer along the southeast than southwest coast of the Izu Peninsula, central Japan. Two bays in this region that have natural populations of E. cava, but with different stipe lengths, were chosen for transplant experiments to examine if stipe length was an environmentally controlled trait. Transplant experiments were carried out in order to determine whether large-type sporophytes of E. cava with long stipes growing in Nabeta Bay (southeast Izu Peninsula, Japan) would turn into small-type sporophytes with short stipes when transplanted to Nakagi Bay (southwest Izu Peninsula). Ten juvenile sporophytes of E. cava (stipe length < 5 cm) were collected from Nabeta Bay (large-type habitat) and transplanted to Nakagi Bay (short-type habitat) in December 1995. As a transplant control, ten juvenile sporophytes of E. cava growing in Nakagi Bay were also transplanted to the same artificial reefs. Growth and survival rates of the sporophytes were monitored monthly for 3 y until December 1998. The transplanted sporophytes showed an increase in their stipe length and diameter from winter to spring, whereas almost no increase was observed from summer to autumn. However, the elongation was greater in Nabeta sporophytes than in Nakagi sporophytes. The primary blade length increased mainly from winter to early spring and decreased largely in autumn. Average primary blade lengths were similar in both Nabeta and Nakagi sporophytes from the end of the first year of transplanting. Although ca. 70% of both Nabeta and Nakagi sporophytes survived during the first 2 y after transplantation, no Nakagi sporophytes and only two Nabeta sporophytes survived to the end of the 3 y study period. Despite transplantation to Nakagi Bay, where short sitpes are naturally present, the sporophytes from Nabeta Bay persisted in having longer stipes, which suggests that stipe length is genetically, rather than environmentally, controlled.     

Cultivation and utilization of Undaria pinnatifida (wakame) as food

A brief review is presented concerning wakame, Undaria pinnatifida, one of the most popular seaweeds used for food in Japan. Although it has been cultivated since about 1940, full-scale cultivation occurred after 1955. As methods for providing ‘seed stock’ and of processing the harvested sporophytes progressed, the yield increased rapidly. The main areas of cultivation are in Japan (e.g. Sanriku, Naruto), Korea and China, while ‘wild’ U. pinnatifida has been introduced into France, New Zealand and Australia. The total world yield of wakame exceeds 500 000 t fresh weight. Cultivated and harvested Undaria is boiled and salted (thus becoming green) and refrigerated; in the factory, it is removed its foreign matter and salt and dried. After checking for quality, the product is packaged in forms convenient for cooking and eating.     

Photosynthesis and respiration in bladelets of Ecklonia cava Kjellman (Laminariales, Phaeophyta) in two localities with different temperature conditions

Characteristics of photosynthesis and respiration of bladelets were compared between Ecklonia cava Kjellman sporophytes growing in a warmer temperate locality (Tei, Kochi Pref., southern Japan) and in a cooler temperate locality (Nabeta, Shizuoka Pref., central Japan). Photosynthesis and respiration were measured with a differential gas-volumeter (Productmeter). In photosynthesis-light curves at 20°C, the rate of net photosynthesis was almost the same at light intensities lower than 25 μmol m⁻² s⁻¹ and the light-saturation occurred at 200–400 μmol m⁻²s⁻¹ in plants of both localities. The light-saturated net photosynthetic rates were higher in winter and spring than in summer and autumn in both plants. The optimum temperature for net photosynthesis at 400 μmol m⁻²s⁻¹ was 27°C throughout the year in the Tei plant and 25–27°C in the Nabeta plant. The decrease of net photosynthetic rates in the supraoptimal temperature range up to 29°C was sharper in winter and spring than in summer and autumn in both plants, being smaller in the Tei plant than in the Nabeta plant in all seasons. The dark respiration rate always increased with water temperature rise in both plants. No clear differences were found in the dark respiration rate between Tei and Nabeta plants except that when measured against dry weight, the Tei plant showed a slightly lower rate as compared with the Nabeta plant.     

The effects of warming on the ecophysiology of two co-existing kelp species with contrasting distributions

The northeast Atlantic has warmed significantly since the early 1980s, leading to shifts in species distributions and changes in the structure and functioning of communities and ecosystems. This study investigated the effects of increased temperature on two co-existing habitat-forming kelps: Laminaria digitata, a northern boreal species, and Laminaria ochroleuca, a southern Lusitanian species, to shed light on mechanisms underpinning responses of trailing and leading edge populations to warming. Kelp sporophytes collected from southwest United Kingdom were maintained under 3 treatments: ambient temperature (12 °C), +3 °C (15 °C) and +6 °C (18 °C) for 16 days. At higher temperatures, L. digitata showed a decline in growth rates and Fv/Fm, an increase in chemical defence production and a decrease in palatability. In contrast, L. ochroleuca demonstrated superior growth and photosynthesis at temperatures higher than current ambient levels, and was more heavily grazed. Whilst the observed decreased palatability of L. digitata held at higher temperatures could reduce top-down pressure on marginal populations, field observations of grazer densities suggest that this may be unimportant within the study system. Overall, our study suggests that shifts in trailing edge populations will be primarily driven by ecophysiological responses to high temperatures experienced during current and predicted thermal maxima, and although compensatory mechanisms may reduce top-down pressure on marginal populations, this is unlikely to be important within the current biogeographical context. Better understanding of the mechanisms underpinning climate-driven range shifts is important for habitat-forming species like kelps, which provide organic matter, create biogenic structure and alter environmental conditions for associated communities.