983 resultados para Pisciculture - Reproductive cycle


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Melatonin regulates the reproductive cycle, energy metabolism and may also act as a potential antioxidant indoleamine. The present study was undertaken to investigate whether long-term melatonin treatment can induce reproductive alterations and if it can protect ovarian tissue against lipid peroxidation during ovulation. Twenty-four adult female Wistar rats, 60 days old (± 250-260 g), were randomly divided into two equal groups. The control group received 0.3 mL 0.9% NaCl + 0.04 mL 95% ethanol as vehicle, and the melatonin-treated group received vehicle + melatonin (100 µg·100 g body weight-1·day-1) both intraperitoneally daily for 60 days. All animals were killed by decapitation during the morning estrus at 4:00 am. Body weight gain and body mass index were reduced by melatonin after 10 days of treatment (P < 0.05). Also, a marked loss of appetite was observed with a fall in food intake, energy intake (melatonin 51.41 ± 1.28 vs control 57.35 ± 1.34 kcal/day) and glucose levels (melatonin 80.3 ± 4.49 vs control 103.5 ± 5.47 mg/dL) towards the end of treatment. Melatonin itself and changes in energy balance promoted reductions in ovarian mass (20.2%) and estrous cycle remained extensive (26.7%), arresting at diestrus. Regarding the oxidative profile, lipid hydroperoxide levels decreased after melatonin treatment (6.9%) and total antioxidant substances were enhanced within the ovaries (23.9%). Additionally, melatonin increased superoxide dismutase (21.3%), catalase (23.6%) and glutathione-reductase (14.8%) activities and the reducing power (10.2% GSH/GSSG ratio). We suggest that melatonin alters ovarian mass and estrous cyclicity and protects the ovaries by increasing superoxide dismutase, catalase and glutathione-reductase activities.

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In the field, mosquitoes characteristically feed on sugars soon after emergence and intermittently during their adult lives. Sugar meals are commonly derived from plant nectar and homopteran honeydew, and without them, adults can only survive for a few days on larval reserves. In addition to sugar, females of most species rely on blood for the initiation and maintenance of egg development; thus their reproductive success depends to some extent on the availability of blood hosts. Males, on the other hand, feed exclusively on sugars. Consequently, their sexual maturation and reproductive success is largely dependent upon access to sugar sources. Plant nectar and homopteran honeydew are the two main sugar sources utilized by mosquitoes in the wild. Previous laboratory studies had shown that differences between nectar sources can affect the survivorship and biting frequency of disease vectoring mosquitoes. However, little is known on how sugar composition influence the reproductive processes in male mosquitoes. Male mosquitoes transfer accessory gland proteins and other hormones to their mates along with sperm during mating. In the female, these seminal fluid constituents exert their influence on reproductive genes that control ovulation and vitellogenesis. The present study tests the hypothesis that the mates of males consuming different sugar meals will exhibit varying levels of induction of vitellogenin (a gene which regulates the expression of egg yolk precursor proteins). Real-time quantitative RT-PCR was used to investigate how each sugar meal indirectly influences vitellogenin mRNA abundance in female Anopheles stephensi following mating. Results indicate that mates of nectar-fed males exhibit 2-fold greater change in vitellogenin expression than the mates of honeydew-fed males. However, this response did not occur in non-blood fed controls. These findings suggest that the stimulatory effect of mating on vitellogenesis in blood meal-reliant (i.e. anautogenous) mosquitoes may only be synergistic in nature. The present study also sought to compare the potential fitness costs of mating incurred by females that do not necessarily require a blood meal to initiate a reproductive cycle (i.e., exhibit autogeny). Females of the facultatively autogenous mosquito, Culex molestus were allowed to mate with males sustained on either nectar or honedyew. Mean lifetime fecundity and survivorship of females under the two different mating regimes were then recorded. Additionally, one-dimensional gel electrophoresis was used to verify the transfer of male accessory gland proteins to the sperm storage organs of females during mating.While there was no significant difference in survival between the test treatments, the mates of nectar-fed males produced 11% more eggs on average than mates of honeydew-fed males. However, additional data are needed to justify the extrapolation of these findings to natural settings. These findings prompt further investigation as the differences caused by diet variation in males may be reflected across other life history traits such as mating frequency and insemination capacity.

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Introduction: L'homéostasie du cholestérol est indispensable à la synthèse de la testostérone dans le tissu interstitiel et la production de gamètes mâles fertiles dans les tubules séminifères. Les facteurs enzymatiques contribuent au maintien de cet équilibre intracellulaire du cholestérol. L'absence d'un ou de plusieurs enzymes telles que la HMG-CoA réductase, la HSL et l'ACAT-1 a été associée à l'infertilité masculine. Toutefois, les facteurs enzymatiques qui contribuent au maintien de l'équilibre intra-tissulaire du cholestérol n'ont pas été étudiés. Cette étude a pour but de tester l'hypothèse que le maintien des taux de cholestérol compatibles avec la spermatogenèse nécessite une coordination de la fonction intracellulaire des enzymes HMG-CoA réductase, ACAT1 et ACAT2 et la HSL. Méthodes: Nous avons analysé l'expression de l’ARNm et de la protéine de ces enzymes dans les fractions enrichies en tubules séminifères (STf) de vison durant le développement postnatal et le cycle reproductif annuel et dans les fractions enrichies en tissu interstitiel (ITf) et de STf durant le développement postnatal chez la souris. Nous avons développé deux nouvelles techniques pour la mesure de l'activité enzymatique de la HMG-CoA réductase et de celle de l'ACAT1 et ACAT2. En outre, l'immunohistochimie a été utilisée pour localiser les enzymes dans le testicule. Enfin, les souris génétiquement déficientes en HSL, en SR-BI et en CD36 ont été utilisées pour élucider la contribution de la HMG-CoA réductase, l'ACAT1 et l'ACAT2 et la HSL à l'homéostasie du cholestérol. Résultats: 1) HMG-CoA réductase: (Vison) La variation du taux d’expression de l’ARNm de la HMG-CoA réductase était corrélée à celle de l'isoforme de 90 kDa de la protéine HMG-CoA réductase durant le développement postnatal et chez l'adulte durant le cycle reproductif saisonnier. L'activité enzymatique de la HMG-CoA réductase augmentait de façon concomitante avec le taux protéinique pour atteindre son niveau le plus élevé à 240 jours (3.6411e-7 mol/min/μg de protéines) au cours du développement et en Février (1.2132e-6 mol/min/μg de protéines) durant le cycle reproductif chez l’adulte. (Souris), Les niveaux d'expression de l'ARNm et l'activité enzymatique de la HMG-CoA réductase étaient maximales à 42 jours. A l'opposé, le taux protéinique diminuait au cours du développement. 2) HSL: (Vison), l'expression de la protéine de 90 kDa de la HSL était élevée à 180- et 240 jours après la naissance, ainsi qu'en Janvier durant le cycle saisonnier chez l'adulte. L'activité enzymatique de la HSL augmentait durant le développement pour atteindre un pic à 270 jours (36,45 nM/min/μg). Chez l'adulte, l'activité enzymatique de la HSL était maximale en Février. (Souris) Le niveau d’expression de l'ARNm de la HSL augmentait significativement à 21-, 28- et 35 jours après la naissance concomitamment avec le taux d'expression protéinique. L'activité enzymatique de la HSL était maximale à 42 jours suivie d'une baisse significative chez l'adulte. 3) ACAT-1 et ACAT-2: Le présent rapport est le premier à identifier l’expression de l'ACAT-1 et de l'ACAT-2 dans les STf de visons et de souris. (Vison) L'activité enzymatique de l'ACAT-2 était maximale à la complétion du développement à 270 jour (1190.00 CPMB/200 μg de protéines) et en janvier (2643 CPMB/200 μg de protéines) chez l'adulte. En revanche, l'activité enzymatique de l'ACAT-1 piquait à 90 jours et en août respectivement durant le développement et chez l'adulte. (Souris) Les niveaux d'expression de l'ARNm et la protéine de l'ACAT-1 diminuait au cours du développement. Le taux de l'ARNm de l'ACAT-2, à l’opposé du taux protéinique, augmentait au cours du développement. L'activité enzymatique de l'ACAT-1 diminuait au cours du développement tandis que celle de l'ACAT-2 augmentait pour atteindre son niveau maximal à 42 jours. 4) Souris HSL-/ -: Le taux d’expression de l'ARNm et l'activité enzymatique de la HMG-CoA réductase diminuaient significativement dans les STf de souris HSL-/- comparés aux souris HSL+/+. Par contre, les taux de l'ARNm et les niveaux des activités enzymatiques de l'ACAT-1 et de l'ACAT-2 étaient significativement plus élevés dans les STf de souris HSL-/- comparés aux souris HSL+/+ 5) Souris SR-BI-/-: L'expression de l'ARNm et l'activité enzymatique de la HMG-CoA réductase et de l'ACAT-1 étaient plus basses dans les STf de souris SR-BI-/- comparées aux souris SR-BI+/+. A l'opposé, le taux d'expression de l'ARNm et l'activité enzymatique de la HSL étaient augmentées chez les souris SR-BI-/- comparées aux souris SR-BI+/+. 6) Souris CD36-/-: L'expression de l'ARNm et l'activité enzymatique de la HMG-CoA réductase et de l'ACAT-2 étaient significativement plus faibles tandis que celles de la HSL et de l'ACAT-1 étaient inchangées dans les STf de souris CD36-/- comparées aux souris CD36+/+. Conclusion: Nos résultats suggèrent que: 1) L'activité enzymatique de la HMG-CoA réductase et de la HSL sont associées à l'activité spermatogénétique et que ces activités ne seraient pas régulées au niveau transcriptionnel. 2) L'ACAT-1 et de l'ACAT-2 sont exprimées dans des cellules différentes au sein des tubules séminifères, suggérant des fonctions distinctes pour ces deux isoformes: l'estérification du cholestérol libre dans les cellules germinales pour l'ACAT-1 et l'efflux du cholestérol en excès dans les cellules de Sertoli au cours de la spermatogenèse pour l'ACAT-2. 3) La suppression génétique de la HSL diminuait la HMG-CoA réductase et augmentait les deux isoformes de l'ACAT, suggérant que ces enzymes jouent un rôle critique dans le métabolisme du cholestérol intratubulaire. 4) La suppression génétique des transporteurs sélectifs de cholestérol SR-BI et CD36 affecte l'expression (ARNm et protéine) et l'activité des enzymes HMG-CoA réductase, HSL, ACAT-1 et ACAT-2, suggérant l'existence d’un effet compensatoire entre facteurs enzymatiques et non-enzymatiques du métabolisme du cholestérol dans les fractions tubulaires. Ensemble, les résultats de notre étude suggèrent que les enzymes impliquées dans la régulation du cholestérol intratubulaire agissent de concert avec les transporteurs sélectifs de cholestérol dans le but de maintenir l'homéostasie du cholestérol intra-tissulaire du testicule.

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The present investigations have considerably enhanced the existing knowledge on the biology and distribution/availability pattern of D.incarnatus in the Malippuram region. The species occurs in good concentration during October - March/April, and disappears from the area during late premonsoon and monsoon months. Recolonising the area in September, it grows fast in the subsequent months. The life span of the species is estimated to be about an year. Studies on the reproductive biology of the species have revealed that there are two spawning peaks, the major peak in February - March and minor peak, in December. The salinity regime of the area influences the reproductive activity. These observations form the original contribution in the thesis. The information on variation in water content, protein,glycogen and lipid levels in relation to reproductive cycle has helped to a better understanding of the gametogenic activity and spawning of the species. Similarly, the findings on salinity tolerance and filtration rate have shown that small sized clams exhibit greater tolerance range than larger clams, and grow at a faster rate with active metabolism. It is hoped that these information would considerably add to the present knowledge of the basic facts which are relevant to the improvement and management of the clam fishery of this region.

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Several studies on the biology and fisheries of mullets, particularly of M cephalus are now available. Different aspects of breeding, larval rearing, seed production, field culture and ecophysiology have also been investigated. However, information on the spermatogenesis in M cephalus as well as L parsia is scanty. Since an understanding of the reproductive strategies is an essential pre-requisite for evolving successful breeding programmes through artificial fertilization and gametic preservation, investigations on spermatogenesis in these species were taken up and the results are presented in this thesis. The thesis is presented in 9 chapters

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Lactation is an energy-demanding process characterized by massive food and water consumption, cessation of the reproductive cycle and induction of maternal behavior. During lactation, melanin-concentrating hormone (MCH) mRNA and peptide expression are increased in the medial preoptic area (MPO) and in the anterior paraventricular nucleus of the hypothalamus. Here we show that MCH neurons in the MPO coexpress the GABA synthesizing enzyme GAD-67 mRNA. We also show that MCH neurons in the MPO of female rats are innervated by neuropeptides that control energy homeostasis including agouti-related protein (AgRP), alpha-melanocyte stimulating hormone (alpha MSH) and cocaine- and amphetamine-regulated transcript (CART). Most of these inputs originate from the arcuate nucleus neurons. Additionally, using injections of retrograde tracers we found that CART neurons in the ventral premammillary nucleus also innervate the MPO. We then assessed the projections of the female MPO using injections of anterograde tracers. The MPO densely innervates hypothalamic nuclei related to reproductive control including the anteroventral periventricular nucleus, the ventrolateral subdivision of the ventromedial nucleus (VMHvl) and the ventral premammillary nucleus (PMV). We found that the density of MCH-ir fibers is increased in the VMHvl and PMV during lactation. Our findings suggest that the expression of MCH in the MPO may be induced by changing levels of neuropeptides involved in metabolic control. These MCH/GABA neurons may, in turn, participate in the suppression of cyclic reproductive function and/or sexual behavior during lactation through projections to reproductive control sites. (C) 2009 Elsevier B.V. All rights reserved.

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Calcium (Ca2+) is a critical regulator of many aspects of the Plasmodium reproductive cycle. In particular, intra-erythrocyte Plasmodium parasites respond to circulating levels of the melatonin in a process mediated partly by intracellular Ca2+. Melatonin promotes the development and synchronicity of parasites, thereby enhancing their spread and worsening the clinical implications. The signalling mechanisms underlying the effects of melatonin are not fully established, although both Ca2+ and cyclic AMP (cAMP) have been implicated. Furthermore, it is not clear whether different strains of Plasmodium use the same, or divergent, signals to control their development. The aim of this study was to explore the signalling mechanisms engaged by melatonin in P. chabaudi, a virulent rodent parasite. Using parasites at the throphozoite stage acutely isolated from mice erythrocytes, we demonstrate that melatonin triggers cAMP production and protein kinase A (PKA) activation. Interestingly, the stimulation of cAMP/PKA signalling by melatonin was dependent on elevation of Ca2+ within the parasite, because buffering Ca2+ changes using the chelator BAPTA prevented cAMP production in response to melatonin. Incubation with melatonin evoked robust Ca2+ signals within the parasite, as did the application of a membrane-permeant analogue of cAMP. Our data suggest that P. chabaudi engages both Ca2+ and cAMP signalling systems when stimulated by melatonin. Furthermore, there is positive feedback between these messengers, because Ca2+ evokes cAMP elevation and vice versa. Melatonin more than doubled the observed extent of parasitemia, and the increase in cAMP concentration and PKA activation was essential for this effect. These data support the possibility to use melatonin antagonists or derivates in therapeutic approach.

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Sea urchins are common benthic organisms on coastal ecosystems of tropical and temperate shallow waters. The impact of sea urchins populations in shore communities is density-dependent, and therefore, knowledge of the life history of these animals is important to understand these interactions. Between 2000 and 2005 a population boom of Tripneustes ventricosus was observed in the Fernando de Noronha Archipelago. In 2004 a research program was started to monitor the population dynamics of T. ventricosus in the archipelago, when it noted a lack of basic information on the biology and reproduction of this species, despite its broad geographic distribution and economic importance in parts of its occurrence. In this context, this work focuses on the reproductive biology of T. ventricosus with emphasis on the description of the gametogenic stages. Between December 2006 and July 2007, ten urchins were collected by snorkeling in two sites of the archipelago, totaling 120 individuals. Gametogenic stages were described for both sexes through microscopic analysis, and were defined as: Recovery, Growing, Premature, Mature and Spawning. Results showed increasing in Gonad index throughout of months of sampling and suggest that the reproductive cycle of the species in the archipelago is annual

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The feeding habits and biological aspects of the reproductive cycle of two-spot astyanax, Astyanax cf. lacustris (Reinhardt, 1874) were investigated. Fish samples were captured on a monthly basis, using gillnets of 4 cm mesh size, from the Piató Lake, Assu, Rio Grande do Norte, during the period of September, 2006 to August, 2007. Physico-chemical parameters, such as, temperature, electrical conductivity and dissolved oxygen of the lake were registered. The monthly values of rainfall also were obtained. The 360 individuals captured, were measured, weighed, dissected, and stomach weight and the stage of gonadal maturity were registered. The stomach contents analyses were carried out based on volumetric method, points, frequency of occurrence and applying the Index of Relative Importance. The degrees of repletion of the stomachs were determined besides the Index of Repletion relating to feeding activity variations and frequency of ingestion with limnological parameters and rainfall. The food items identified were separated into distinct groups according to their origin. Sex ratio and Gonadosomatic Relation of females were verified to determine the spawning period and fecundity. The physico-chemical parameters presented the following annual mean values: temperature = 28.8ºC, electrical conductivity = 992.5 µS.cm-1; dissolved oxygen = 4.9 mg.L-1 during the study period. The annual mean of the rainfall was 63.5 mm. The results indicate that this species present an omnivorous feeding habit with a tendency towards insectivory, with an increase in feeding activity during the dry season. The aquatic oxygen to interfere very importance in the feeding activity than the others factors physico-chemicals of water and rainfall. There was a predominance of females, with a sex proportion of 1M:7F. The macroscopic characteristics of the ovaries and testicles revealed four stages of gonadal development: immature, maturing, mature and spent. A temporal variation was observed for the gonadal development of males and females. There was reproductive activity through out the year, with peaks in the months of February, April and June to correspond with the rain of precipitation of the region. The mean fecundity was 7.681 mature oocytes, varying from 4.476 to 12.036, with mean of 7.681. There was positive relation between fecundity and body mass. Condition Factor is not an efficient indicator of the reproductive period of this species. The species A. cf. lacustris is an opportunist and is well adapted to the conditions of the semi-arid Caatinga Biome

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The reproductive ecology of fish plays a key role both for rational exploitation methods and for protective measures of natural stocks. The purpose of this study was to analyze the reproductive aspects of the damsel-fish, Stegastes fuscus, during October 2004 to September 2005, in the coastal rocky reefs of Búzios Beach, Nísia Floresta, RN. Fish were captured using hooks and hand nets, during low tide. Reproduction was determined using sexual ratio, mean length of first maturation (L50), absolute fecundity and macroscopic characteristics of gonads. The following parameters were related to gonadosomatic index (GSI): condition factor (CF), hepatosomatic index (HSI), rain fall and temperature. In relation to sex distribution, it was observed that 78% were females and 22% were males. The L50 was 6.2 cm for females and 7.0 for males. Average fecundity was 6832 oocytes. Results showed that S. fuscus had better body condition in the months prior to spawning, particularly during initial and intermediate stages of maturation. Five stages of gonadal maturation were identified through macroscopic analysis: immature, in maturation, mature, spent and resting. The HSI was inversely related to the GSI. This was possibly due to the reproductive cycle of this species which was associated to the dry period of this region. During this period, low rain fall and high temperatures provide an propitious reproductive condition for the study species

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In the months of January 2001 and 2002, female cachara Pseudoplatystoma fasciatum were selected during their first and second gonadal maturation (2 years and 7 months old and 3 years and 7 months old, respectively) with an of oocyte diameter of 937.5 mum (82.5% with central nuclei and 17.5% with peripheral nuclei). Nine females in first maturation received two doses of carp pituitary extract (CPE), 0.5 mg/kg and 5.0 mg/kg; seven received two doses of human chorionic gonadotropin (hCG), 5 and 10 IU/g; five received doses of 0.5 CPE mg/kg and 5 hCG IU/g (CPE+hCG); and four received 0.9% saline (saline). Nine females from CPE and seven from hCG presented oocytes with the same diameter at the moment of oocyte release (100% with germinal vesicle breakdown and fertilization rate of 53.44 +/- 18.3 and 54.81 +/- 11.8%; larvae number of 165,330 +/- 94.1 and 158,570 +/- 20.6, respectively). The five females from CPE+hCG did not respond to the hormonal treatment. The four females from the saline group did not ovulate. In January 2002, 6 of 15 selected females that were going through the second reproductive cycle received CPE (five received hCG and four received saline), showing oocyte diameters similar to the ones in the first maturation. At stripping, CPE females had an oocyte diameter of 1062.5 mum (the hCG females had oocyte diameters ranging from 937.5 to 1125.0 mum; fertilization rates of 56.08 +/- 30.9 and 81.90 +/- 17.3%; 364,547 +/- 244 and 633,129 +/- 190, larvae, respectively). The fertilization rates and larvae number were higher in the second gonad maturation, both for CPE and hCG. (C) 2004 Elsevier B.V. All rights reserved.

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The eutrophication (cultural or anthropogenic) is induced by man and can have different origins, such as domestic sewage, industrial and agricultural activities, including the effluent still breeding systems of aquatic organisms. The expansion of aquaculture, with production of biomass and increase of nutrients in water may cause acceleration of productivity of algae, changing the ecology of aquatic systems. In addition, these waste water may present a risk to health through the transfer of pathogens from manure, plant residues, composted material, among others that are major sources of organic waste in some farming systems. Depending on the trophic level of fish ponds, which are dynamic environments, different planktonic species with short reproductive cycle and adapted to the changes contained in these systems can appear in high abundance. Water quality in the systems for raising fish is related to several factors, such as water source, management (liming, fertilizing, cleaning), cultivated species and quantity and composition of exogenous food. In order to minimize environmental impacts, there are techniques to improve the quality of water in fish farming systems and thus satisfactory answers can be obtained through the application of management practices. This paper aims to review the subject that deals with changes in water quality resulting from the activity of freshwater fish culture in Brazil. Search also recommend techniques of good management practices to minimize the impact generated by the activity.

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For a long time, we believed in the pattern that tropical and south hemisphere species have high survival. Nowadays results began to contradict this pattern, indicating the need for further studies. Despite the advanced state of the study of bird population parameters, little is known about their variation throughout the year and the factors affecting them. Reproduction, for example, is one factor that may alter adult survival rates, because during this process the breeding pair allocates resources to maintain itself to maintain offspring, making itself more susceptible to diseases and predation. The aim of this study was to estimate survival and population size of a Central and South America passerine, Tachyphonus rufus (Boddaert, 1783), testing hypotheses about the factors that define these parameters. We performed data collection between Nov/2010 and ago/2012 in 12 ha plot, in a fragment of Atlantic Forest in northeastern Brazil. We used capture-mark-recapture methods to generate estimates using Closed Design Robust model in the program MARK. We generated Multi-state models to test some assumptions inherent to Closed Robust Design. The influence of co-variables (time, rain and reproductive cycle) and the effect of transient individuals were measured. Capture, recapture and apparent survival parameters were defined by reproductive cycle, while temporary dispersal was influence by rain. The estimates showed a higher apparent survival during the non-breeding period (92% ± 1%) than during breeding (40% ± 9%), revealing a cost of reproduction and suggesting a trade-off between surviving and reproducing. The low annual survival observed (34%) did not corroborate the pattern of high rates expected for a tropical bird. The largest population size was estimated to be 56 individuals in Nov/11, explained by high recruitment of juveniles, while the lowest observed in May/12: 10 individuals, probably as a result of massive influx of competitor species. Results from this study add to the growing literature on life history of Neotropical species. We encourage studies like this especially in Brazil, where there are few information, and suggest that covariates related to habitat quality and environmental changes should be tested, so that we can generate increasingly reliable models

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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

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O presente estudo teve por objetivo avaliar o desenvolvimento de frutos de plantas de longan (Dimocarpus longan Lour) na região de Jaboticabal, estabelecendo-se, portanto, a curva de crescimento dos frutos. As avaliações realizadas foram semanais, consistindo em medições do diâmetro longitudinal e transversal dos frutos marcados, a partir do momento da formação da polpa. Quinzenalmente, realizaram-se avaliações de características físicas: massa de fruto (g), casca (g), polpa (g) semente (g), e químicas: acidez titulável (AT), sólidos solúveis (SS), ácido ascórbico (AA) e ratio (SS/AT). Com os resultados obtidos, pode-se concluir que o ciclo de frutificação da Longan na região é entre 120 e 130 dias após antese, período em que o fruto estaria apto para o consumo. A curva do desenvolvimento dos frutos segue um padrão sigmoidal simples. Com relação aos atributos físicos e químicos, constatou-se que a planta 6 apresentou os melhores resultados, podendo ser recomendada como planta-matriz.