Neural Correlates of Migration: Activation of Hypothalamic Clock(s) in and out of Migratory State in the Blackheaded Bunting (Emberiza melanocephala)

Background

Many vertebrates distinguish between short and long day lengths using suprachiasmatic nuclei (SCN). In birds particular, the mediobasal hypothalamus (MBH) is suggested to be involved in the timing of seasonal reproduction. This study investigated the response of SCN and MBH to a single long day, and the role of MBH in induction of the migratory phenotype in night-migratory blackheaded buntings.

Methodology/Principal Findings

Experiment 1 immunocytochemically measured c-fos in the SCN, and c-fos, vasoactive intestinal peptide (VIP) and neuropeptide Y (NPY) in the MBH of buntings exposed to a 20 h light period. Long light period induced significantly stronger c-fos expression, measured as number of c-fos-like immunoreactive (c-fos-lir) cells, in MBH, but not in the SCN. Within the MBH, c-fos-lir cells were significantly denser in the inferior hypothalamic nucleus (IH) and infundibular nucleus (IN), but not in the dorsomedial hypothalamus (DMH). IH and IN also had significantly increased number of VIP and NPY labeled cells. DMH had significantly increased number of VIP labeled cells only. Experiment 2 assayed c-fos, VIP and NPY immunoreactivities in the middle of day and night in the MBH of buntings, after seven long days (day active, non-migratory state) and after seven days of Zugunruhe (night active, migratory state) in long days. In the migratory state, the number of c-fos-lir cells was significantly greater only in DMH; VIP-lir cells were denser in all three MBH regions suggesting enhanced light sensitivity at night. The denser NPY-lir cells only in IN in the non-migratory state were probably due to premigratory hyperphagia.

Conclusions/Significance

In buntings, SCN may not be involved in the photoperiod-induced seasonal responses. MBH contains the seasonal clock sensitive to day length. VIP and NPY are parts of the neuroendocrine mechanism(s) involved, respectively, in sensing and translating the photoperiodic message in a seasonal response.     

Seasonal modulation of diurnal food consumption in Indian songbirds

Seasonal changes in daily food consumption have a direct bearing with energy requirement of bird that is in turn associated with life history stage of birds. We compared seasonal changes in daily food intake in adult male migratory redheaded bunting (Emberiza bruniceps) that over winters in Indian subcontinent with those in non-migratory blackheaded munia to reiterate the same. We also compared daily food eating pattern (DFEP) in wintering blackheaded and redheaded buntings, closely related Emberizidae finches to establish circadian nature of feeding behavior and how it varied at species level. The birds were held under short days (8L:16D; 8 h of light and 16 h of darkness) and two hourly food consumption was measured to profile their DFEP. Further, we extended the study to establish how the circadian pattern of food consumption varied depending on birds’ physiological state and effect of photoperiod in adult male redheaded buntings. Redheaded buntings DFEP and locomotor activity were compared in pre-migratory months of February (spring) and September (autumn). The results suggest that September (photorefractory) birds exhibit clear bimodality in their feeding behavior as compared to (photosensitive) birds in February. Another experiment compared bird’s DFEP held under short (8L:16D) and long (16L:8D) days for 5 weeks and suggested that under long days, prolonged hours of photophase render adaptive advantage to birds for positive energy budgeting. The present study clearly establishes the circadian nature of feeding behavior and that it modulates over seasons. The bimodal i.e. morning and evening peaks of food consumption suggest morning–evening food entrainable oscillators, however this needs to be investigated with mechanistic approach in future studies.     

The tempo-spatial variations of phytoplankton diversities and their correlation with trophic state levels in a large eutrophic Chinese lake

Lake Chaohu is one of the most eutrophic lakes in China. Research on this lake's seasonal and spatial variations in phytoplankton diversity is needed to understand the distribution of eutrophication, as well as to find appropriate comprehensive biodiversity indices to assess the eutrophication status of the lake. The present study indicated that the Margalef index of all samples was as low as 0.799 ± 0.543 in summer (August 2011) and as high as 1.467 ± 0.653 in winter (February 2012). The Margalef index of the river samples had a high mean value and substantial variation compared with the lake samples. The Peilou index of the lake samples was higher than that of the river samples in summer and autumn (November 2011) but lower than that of the river samples in winter. In spring (May 2012), the Peilou index of the western samples was lower than that of the eastern samples. The spatial distribution of the Shannon–Wiener index was more similar to that of the Peilou index in autumn and winter, while in spring and summer, the spatial distribution was affected by both species richness and evenness. High eutrophication levels occurred in the western lake in spring and summer, whereas high levels occurred in the eastern lake, especially in the middle of the lake, in autumn and winter. The total trophic state index (TSI) in all samples exhibited a significant negative correlation with the Margalef (r = −0.726) and Peilou (r = −0.530) indices but a significant positive correlation with the Shannon–Wiener (r = 0.654) index. The partial correlation analysis results implied that these phytoplankton biodiversity indices could serve as synthetic ecological indicators to assess the eutrophication condition of Lake Chaohu.     

Investigations of photoperiodically induced fattening in migratory blackheaded bunting (Emberiza melanocephala)(Ayes)

The body fattening and weight gain preceding vernal migration in birds is timed by a set of environmental factors of which daylength is predominant. However, the mechanism(s) by which these events is determined is poorly understood. Previous investigations on a photoperiodic migratory species, the blackheaded bunting ( Emberiza melanocephala ), indicate the involvement of a light-sensitive circadian rhythm during initiation of fat deposition and body weight gain. This communication presents data from another set of experiments aimed to characterize further the mechanism(s) of fat deposition in the same species.
Groups of photosensitive, unstimulated and stimulated birds were subjected to transfer and superimposition experiments for 30 days. While the former set included shifting of long-day (LD) birds to DD, SD (short days), DD/LD and SD/LD, in the latter a 90-minute bright light was superimposed at two different times of the day during the dim-green lighted phase 15L:9D of varying intensity. Birds were weighed at the beginning and at the end of experiments. Those in transfer cycles were also weighed at 10-day intervals. The results suggest that the premigratory body fattening and weight gain in blackheaded buntings is light dependent and timed by environmental daylength in accordance with the photosensitive endogenous circadian rhythm (ECR). They also show that the photoperiodic responses in birds in general are mediated by circadian rhythm(s).     

Temperature alters the photoperiodically controlled phenologies linked with migration and reproduction in a night-migratory songbird

We investigated the effects of temperature on photoperiodic induction of the phenologies linked with migration (body fattening and premigratory night-time restlessness, Zugunruhe) and reproduction (testicular maturation) in the migratory blackheaded bunting. Birds were exposed for four weeks to near-threshold photoperiods required to induce testicular growth (11.5 L:12.5 D and 12 L:12 D) or for 18 weeks to a long photoperiod (13 L:11 D) at 22°C or 27°C (low) and 35°C or 40°C (high) temperatures. A significant body fattening and half-maximal testicular growth occurred in birds under the 12 L, but not under the 11.5 L photoperiod. Further, one of six birds in both temperature groups on 11.5 L, and four and two of six birds, respectively, in low- and high-temperature groups on 12 L showed the Zugunruhe. Buntings on 13 L in both temperature groups showed complete growth-regression cycles in body fattening, Zugunruhe and testis maturation. In birds on 13 L, high temperature attenuated activity levels, delayed onset of Zugunruhe by about 12 days, reduced body fattening and slowed testicular maturation. The effect of temperature seems to be on the rate of photoperiodic induction rather than on the critical day length. It is suggested that a change in temperature could alter the timing of the development of phenologies linked with seasonal migration and reproduction in migratory songbirds.     

Stress impairs new but not established relationships in seasonally social voles

Affiliative social relationships are impacted by stressors and can shape responses to stress. However, the effects of stress on social relationships in different contexts are not well understood. Meadow voles provide an opportunity to study these effects on peer relationships outside of a reproductive context. In winter months, female meadow voles cohabit with peers of both sexes, and social huddling is facilitated by exposure to short, winter-like day lengths in the lab. We investigated the role of stress and corticosterone (cort) levels in social behavior in short day-housed female meadow voles. A brief forced swim elevated cort levels, and we assessed the effects of this stressor on new and established relationships between females. In pairs formed following exposure to swim stress, the stressor significantly reduced the fraction of huddling time subjects spent with a familiar partner. Swim stress did not affect partner preferences in pairs established prior to the stressor. Finally, we examined fecal glucocorticoid metabolite levels via immunoassay in voles housed under short day (10 h light) versus long day (14 h light) conditions and detected higher glucocorticoid levels in long day-housed voles. These findings support a role for stress regulation in the formation of social relationships in female meadow voles, and are consistent with a potential role for seasonal variation in cort in the behavioral transition from solitary to social. Together they highlight the importance of stress and possibly glucocorticoid signaling for social behavior.     

Melatonin modulates glucose homeostasis during winter dormancy in a vespertilionid bat,Scotophilus heathi

The role for melatonin in glucose homeostasis and insulin resistance is not very clear and has recently been an active area of investigation. The present study investigated the role of melatonin in seasonal accumulation of adipose tissue in Scotophilus heathi, with particular reference to its role in glucose homeostasis and development of insulin resistance. The circulating melatonin levels correlated positively (p < 0.05) with the changes in body mass due to fat accumulation and circulating insulin level, but correlated negatively with the blood glucose level in S. heathi. The bats showed high circulating blood glucose levels and impaired glucose tolerance during the period of fat deposition suggesting insulin resistance condition which improves after winter when most of the fat has been utilized as a metabolic fuel. The high circulating melatonin levels during the period of maximum body fat at the beginning of winter prepare the bats for winter dormancy by modulating the glucose homeostasis through affecting blood glucose levels, muscle and liver glycogen stores, insulin receptor and glucose transporter 4 (GLUT 4) expression. This is also confirmed by in vivo study in which melatonin injection improves the glucose tolerance, increases muscle insulin receptor and GLUT 4 expression, and enhances glucose clearance from the blood. The results of present study further showed that the effect of melatonin injection on the blood glucose levels is determined by the metabolic state of the bats and may protect from decrease in blood glucose level during extreme starvation, however, melatonin when injected during fed state increases glucose clearance from the blood. In summary, the present study suggested that melatonin interferes with the glucose homeostasis through modulating intracellular glucose transport and may protect bats from hypoglycemia during winter dormancy.     

Body-size spectra of biofilm-dwelling protozoa and their seasonal shift in coastal ecosystems

Community-based assessment of protozoa is usually performed at a taxon-dependent resolution. As an inherent ‘taxon-free’ trait, however, body-size spectrum has proved to be a highly informative indicator to summarize the functional structure of a community in both community research and monitoring programs in aquatic ecosystems. To demonstrate the relationships between the taxon-free resolution of protozoan communities and water conditions, the body-size spectra of biofilm-dwelling protozoa and their seasonal shift and environmental drivers were explored based on an annual dataset collected monthly from coastal waters of the Yellow Sea, northern China. Body sizes were calculated in equivalent spherical diameter (ESD). Among a total of 8 body-size ranks, S2 (19–27 μm), S3 (28–36 μm), S4 (37–50 μm) and S5 (53–71 μm) were the top four levels in frequency of occurrence, while rank S1 (13–17 μm), S2 and S4 were the dominant levels in abundance. These dominants showed a clear seasonal succession: S2/S4 (spring) → S2/S4 (summer) → S4 (autumn) → S2 (winter) in frequency of occurrence; S1 (spring) → S4 (summer) → S2 (autumn) → S1 (winter) in abundance. Bootstrapped average analysis showed a clear seasonal shift in body-size spectra of the protozoa during a 1-year cycle, and the best-matching analysis demonstrated that the temporal variations in frequency of occurrence and abundance were significantly correlated with water temperature, pH, dissolved oxygen (DO), alone or in combination with chemical oxygen demand (COD) and nutrients. Thus, the body-size spectra of biofilm-dwelling protozoa were seasonally shaped and might be used as a time and cost efficient bioindicator of water quality in marine ecosystems.     

Changes in food intake,body weight,gonads and plasma concentrations of thyroxine,luteinizing hormone and testosterone in captive male buntings exposed to natural daylengths at 29° N

We have investigated the seasonal changes in food intake, body weight, gonadal volume and plasma concentrations of thyroxine, luteinizing hormone and testosterone in male blackheaded bunting (Emberiza melanocephala) in captivity under natural daylengths at 29° N. The cycles in food intake, body weight and testis size in buntings appeared to be phase related. While the changes in body weight and testicular size were parallel to each other and correspond to the increasing daylengths of spring and early summer, cycle in food intake was almost antiphase to the cycles in body weight and testicular growth and development. Furthermore, buntings showed a distinct seasonal cycle in plasma concentrations of thyroxine, luteinizing hormone and testosterone. It is suggested that these seasonal cycles in buntings are endogenously programmed and their entrainment to the environmental photoperiod ensures the occurrence of different physiological functions at temporally fixed time of the year.     

Biochronometry of photoperiodically induced fat deposition in a migratory finch, the Blackheaded bunting (Emberiza melanocephala) (Aves)

This investigation attempts to identify the mechanism(s) involved in the fat deposition in a photoperiodic migratory species, the Blackheaded bunting ( Emberiza melanocephala ). Groups of photosensitive male buntings were exposed to resonance, interrupted night, and ultra-short-day light cycles for 35, 42 and 75 days, respectively. Birds were weighed at the beginning and at the end of the experiments. Those exposed to ultra-short-day light cycles were also weighed at critical intervals during the treatment period. Our results indicate that: (a) a light-sensitive rhythm with a period of about 24 hr is involved in the photoperiodic induction of premigratory fattening and weight gain in Blackheaded buntings: (b) buntings possess a bimodal pattern 'of sensitivity to photoperiods that induce fattening, and (c) this endogenous circadian rhythm can be entrained by an ultrashort photophase of 3 h if the latter is coupled with scotophases of specific duration.     

Seasonal changes in the ranging area of Brown-eared pheasant and its affecting factors in Huanglong Mountains, Shaanxi Province

A study on the seasonal changes in the ranging area of Brown-eared pheasant and its affecting factors was conducted in the Huanglongshan Nature Reserve, Huanglong County, Shaanxi Province, China, from March 2006 to January 2007. Forty used sites and forty random plots were selected along 8–9 transects crossing the study area. Fifteen factors related to the changes of altitude were measured in each site. The results showed that the altitude of home range of Brown-eared pheasant varied with seasons: the highest home range was found in summer, and the lowest one was found in winter, while the home range in autumn was higher than that in spring. According to the frequency of occurrences of Brown-eared pheasant in different seasons, we found that its home range occurred mainly at an altitude below 1400 m in spring, above 1500 m in summer, between 1200 m and 1500 m in autumn and below 1300 m in winter. The possible reason that Brown-eared pheasants preferred to live at lower altitude in spring was to have access to water sources and the abundance of food, correlated to the slope locations, number of trees and number of shrubs; in summer, they tended to appear at highest altitude, and the average height of shrubs, average height of grasses, sheltering class and distance to edge of woods, and human disturbance were affecting factors; in autumn, they appeared at higher altitude which was correlated with the covers of trees, shrubs and grasses, and ultimately related to the abundance of food; in winter, they lived at lowest altitude, which was correlated to the distance from the edge of woods, cover of trees and sheltering class, and ultimately related to the abundance of food and shelter.     

Seasonal and annual performance of a full-scale constructed wetland system for sewage treatment in China

A full-scale constructed wetlands system with a total area of 80 ha and treatment capability of 2.0 × 10⁴ m³ d⁻¹ was completed in October 1998 in Rongcheng, Shandong Province, China. To evaluate wastewater treatment effectiveness and seasonal performance of the system, water samples were collected and analyzed from January 1999 to December 2004. Comparison of mean inlet and outlet concentrations showed that the constructed wetland system could effectively reduce the output of SS (71.8 ± 8.4%), BOD₅ (70.4 ± 9.6%), COD (62.2 ± 10.1%), total coliform (99.7%) and fecal coliform (99.6%). However, the percent reduction of ammonia nitrogen was relatively low (40.6 ± 15.3%), and total phosphorus showed the least efficient reduction (29.6 ± 12.8%). BOD₅, COD, ammonia nitrogen, and total phosphorus removal efficiencies displayed seasonal variations. BOD₅ and COD removal was more efficient in spring and summer than in autumn and winter whereas ammonia nitrogen and total phosphorus removal was more efficient in summer and autumn than in spring and winter. Annual variation analysis shows that COD, BOD₅, and ammonia nitrogen reduction efficiencies increased from 1999 to 2004. In contrast, mean total phosphorus reduction efficiency did not change from 2001 to 2002 and began to decrease from 2003 onwards.     

Factors controlling Zostera marina L. growth in the eastern and western Pacific Ocean: Comparisons between Korea and Oregon,USA

Zostera marina distribution is circum-global and tolerates a wide range of environmental conditions. Consequently, it is likely that populations have adapted to local environmental conditions of light, temperature and nutrient supply. We compared Z. marina growth dynamics over a 2-year period in relation to environmental characters at Jindong Bay, South Korea and Yaquina Bay, Oregon, USA. Water temperature in Jindong Bay showed stronger seasonal variation (summer–winter ΔT = 20 °C) than in Yaquina Bay (summer–winter ΔT < 5 °C). Underwater irradiance in Jindong Bay exhibited a winter maximum, while in Yaquina Bay underwater light exhibited a summer maximum. Integrated annual underwater irradiance during 2003 was 2200 and 1200 mol photons m⁻² year⁻¹ in Korea and Oregon, respectively. Z. marina shoot density, biomass and integrated production were not significantly different between the two study sites. Seasonal Z. marina growth in Jindong Bay appeared to be controlled by temperature and light, while the growth pattern in Yaquina Bay suggested light regulation. Several seagrass parameters were correlated to phosphate concentrations, even though nutrients did not appear limiting. Despite differences in environmental factors, relative growth rates and temporal growth dynamics between study sites, integrated annual leaf production was quite similar at 335 and 353 g DW m⁻² year⁻¹ in the Jindong and Yaquina Bay study sites. We suggest that Z. marina net productivity is acclimated to the local environmental conditions and may be a general characteristic of temperate seagrass populations.     

Coexistence of insulin resistance and increased glucose tolerance in pregnant rats: a physiological mechanism for glucose maintenance

AimThe contribution of insulin resistance (IR) and glucose tolerance to the maintenance of blood glucose levels in non diabetic pregnant Wistar rats (PWR) was investigated.Main methodsPWR were submitted to conventional insulin tolerance test (ITT) and glucose tolerance test (GTT) using blood sample collected 0, 10 and 60 min after intraperitoneal insulin (1 U/kg) or oral (gavage) glucose (1 g/kg) administration. Moreover, ITT, GTT and the kinetics of glucose concentration changes in the fed and fasted states were evaluated with a real-time continuous glucose monitoring system (RT-CGMS) technique. Furthermore, the contribution of the liver glucose production was investigated.Key findingsConventional ITT and GTT at 0, 7, 14 and 20 days of pregnancy revealed increased IR and glucose tolerance after 20 days of pregnancy. Thus, this period of pregnancy was used to investigate the kinetics of glucose changes with the RT-CGMS technique. PWR (day 20) exhibited a lower (p < 0.05) glucose concentration in the fed state. In addition, we observed IR and increased glucose tolerance in the fed state (PWR-day 20 vs. day 0). Furthermore, our data from glycogenolysis and gluconeogenesis suggested that the liver glucose production did not contribute to these changes in insulin sensitivity and/or glucose tolerance during late pregnancy.SignificanceIn contrast to the general view that IR is a pathological process associated with gestational diabetes, a certain degree of IR may represent an important physiological mechanism for blood glucose maintenance during fasting.     

Responses of wolf feeding habits after adverse climatic events in central-western Belarus

Many studies have investigated the ecology of wolf populations of Eurasia, showing that although wolves are mostly opportunistic in seeking meso-large enough mammalian prey, they can also be selective, depending on local availability of prey and their population biomass. Yet prey preferences of the wolf have been poorly evaluated in situations of complex predator/prey systems because such ecological situations are extremely rare in Europe. In particular, the role of beaver is poorly known due to the extreme decline in its range and population over the last few centuries.We conducted a 15-year study (1999–2014) of wolf Canis lupus diet in the Naliboki forest of central-western Belarus to determine the dietary responses of the wolf population in a context of a rich prey supply (beaver 650 inds/100 km², elk 47 inds/100 km², red deer 98 inds/100 km, roe deer 398 inds/100 km², wild boar 234 inds/100 km²). The bison, although present, is not preyed on. We compared the seasonal and annual diet variations of both wolf adults and pups, by scat analysis and hair identification. In winter 2012–2013, the winter was quite harsh with a long period of snow, which severely affected the roe deer and wild boar populations. Five severe summer droughts also occurred (1999, 2001, 2002, 2004 and 2013), greatly decreasing the water level in rivers and canals. We took advantage of these stressful events to evaluate the diet responses of the wolves.In “normal” years, we identified 11 food categories, essentially beaver and medium-sized ungulates (66%), and large ungulates to a lesser extent (9% in summer, 20% in winter). The adults were found to selectively supply pups with beaver, probably because of its easy transportability. Beaver consumption also increased during summer droughts when water levels were very low. After the harsh winter of 2012–2013, which was followed by a sharp decline in medium-sized prey, we observed a shift in the winter diet breadth of the wolves towards greater consumption of both large wild ungulates and small carnivores. We concluded that:1. Beaver is a functional element in wolf ecology, as a primary food for adults and pups;2. A large range of available prey species is important to maintaining a viable wolf population in cases of extreme climatic events.     

Vitamin D Status and Its Seasonal Variability in Healthy Young Adults in an Asian Indian Urban Population

ObjectiveTo assess the vitamin D status and its seasonal variability in healthy young adults in Chandigarh, a city in northern India.MethodsThe history was elicited and examination was performed pertaining to metabolic bone disease in 329 young adults (18 to 25 years of age) at the end of summer and 237 subjects from the same cohort at the end of winter. The calcium profile, 25-hydroxyvitamin D [25(OH)D], and intact parathyroid hormone were measured during both the seasons.ResultsAmong the young adults in this study cohort, 25(OH)D sufficiency (≥ 30 ng/mL) was found in 72.5% in summer and in 50.7% in winter. A significantly higher number of men were 25(OH)D sufficient in comparison with women in summer (P = .001). The mean (standard deviation) serum 25(OH)D levels were significantly higher at the end of summer—52.9 (33.7) ng/mL—in comparison with that at the end of winter—31.8 (21.1) ng/mL; P < .001. The intact parathyroid hormone levels were significantly lower in the 25(OH)D-sufficient group (P = .001) and began to increase at 25(OH)D levels below 25 ng/mL. The serum 25(OH)D levels correlated positively with the duration of exposure to sunlight during summer (r = 0.111; P = .05) and also the calcium intake during summer (r = 0.129; P = .03).ConclusionVitamin D sufficiency may be a reality with a combination of young skin, optimal and effective exposure to sunlight, and adequate calcium intake. (Endocr Pract. 2011;17:185-191)     

Diet and nutrition of range goats on a sarcocaulescent shrubland from Baja California Sur,Mexico

Botanical composition of diets of range goats was studied to determine seasonal preference indices of forage species and nutritional quality of selected diets on a sarcocaulescent shrubland from the Sonoran desert in Baja California Sur, Mexico. Extrusas from five esophageal cannulated adult female goats (40 kg of BW) were collected at the beginning and at the end of each season from April of 2001 to March of 2002. At the end of spring, goats selected higher amounts of browse legumes and cacti species, but during late summer, autumn and winter goats mainly preferred browse non-legumes followed by forb species. Goats selected a constant diet in truly digestible crude protein (annual mean = 11 ± 0.4% DM) and metabolizable energy (2.4 ± 0.1 Mcal kg⁻¹) throughout the year. However, during late spring and early summer, goats selected diets higher in non-structural carbohydrates (NSC), truly IVDOM and truly digestible NSC. Dietary Ca, Mg, K, Mn, Fe were in substantial amounts to meet requirements of a range goat weighing 40.0 kg BW consuming 76.3 g DM kg^0.75 d⁻¹ in all seasons. It is concluded that range goats may require supplementary protein and energy to overcome pregnancy and lactation requirements throughout the year. Copper content appeared marginally deficient almost the entire year and Zn during late summer and autumn.     

Seasonal changes of mechanisms maintaining clear water in a shallow lake with abundant Chara vegetation

The study is based on monitoring data on the seasonal variation during four (1996–1999) vegetation periods, as well as long-term summer data on submerged vegetation, nutrients, light, phytoplankton and zooplankton in Lake Krankesjön, a shallow, calcium-rich, moderately eutrophic lake in southern Sweden.The lake has been in the clear water state with abundant submerged vegetation since the end of the 1980s. Somewhat lower summer biomass of submerged macrophytes during 1997 and 1999 indicates a temporary instability of the clear water state. During these 2 years, summer transparency was about 1.2–2.1 m, while concentrations of total phosphorus and chlorophyll (Chl) a were about 26–40 and 8–18 μg l⁻¹, respectively.Summer biomass of submerged macrophytes was higher during 1996 and 1998. In both years, a distinct increase in light availability and decrease in concentrations of nutrients and chlorophyll occurred simultaneously with the development of dense Chara vegetation. Summer values for transparency were about 2.0–2.5 m, while concentrations of total phosphorus and Chl a were about 20–32 and 4–11 μg l⁻¹, respectively.Summer biomass of crustacean zooplankton was below 250 μg l⁻¹ during all 4 years. A peak abundance of Cladocera (mainly Bosmina longirostris) during May or June caused only a short-term reduction in chlorophyll concentrations that was more pronounced in 1997 than in 1996.Measured light attenuation during 1999 was closely correlated with light attenuation calculated from the amount of suspended solids, chlorophyll concentrations, and water colour. Detritus contributed most to the total amount of suspended solids, while chlorophyll was the main contributor to light attenuation.A long-term decrease of the ratios between chlorophyll and total phosphorus suggests that phytoplankton in the clear water state is limited by factors other than total phosphorus concentrations. Increased sedimentation rate, carbon limitation, allelopathy and a lower bioavailable fraction of the total amount of phosphorus are possible explanations, while nitrogen limitation and grazing from zooplankton probably are of minor importance.Possible reasons for the “instability” of the clear water state during 1997 and 1999 are discussed. Unusually high water level as well as cold and windy weather during the spring of 1996–1999 may have caused a slow and late growth of the plants and thus a temporary instability. However, a tendency for an increase in total phosphorus concentrations and sediment accumulation along the wind-protected shores during the clear water state indicate the possibility of a long-term destabilization which contradicts the alternative stable states model.     

Vegetation patterns and spontaneous regression of Caulerpa taxifolia (Vahl) C. Agardh in Malinska (Northern Adriatic,Croatia)

The green alga Caulerpa taxifolia was recorded at Malinska in 1994 and this actually represents the highest northern latitude (45°7′30″N) at which this invasive alga has been found in the world. The alga was widespread at four sites from which it was eradicated by suction pumps during 1996 and 1997. However, it immediately and intensively recolonized all but one site. Throughout 1998 and 1999 the renewed vegetation showed consistent seasonal patterns. The alga nearly disappeared in April and May while regenerating from over-wintering parts of the thalli in summer. The maximum development occurred in autumn and winter with values of biomass (around 200 g dry weight m⁻²) and frond number (around 2000 m⁻²) generally lower than those reported for the north-western Mediterranean. Values for the frond length (10–18 cm) were in the same range as those in the north-western basin. Throughout 1998 and 1999 the biomass was closely correlated to frond number and length (adjusted R² = 0.90). During the following years C. taxifolia entered a phase of regression. The total colonized area, which amounted to several thousands of square metres in 1998, spontaneously declined in 2000 and 2001 so that only several thalli were found in 2004 during a detailed survey of the settlement. No major changes in winter seawater temperatures, ranging from 9.5 to 10.5 °C, were observed in the area from 1994 to 2004. Thus, other unknown processes could likely play a role on specific vegetation patterns of C. taxifolia in Malinska. Accordingly, it is difficult to explain why the still surviving thalli did not proliferate during the favourable summer–autumn period.