For everything a season: Smoke-induced seed germination and seedling recruitment in a Western Australian Banksia woodland

Abstract The influence of factors associated with fire on seed germination of Australian native species is generally well documented, but examples involving the use of smoke as a fire analogue for ecological research remain limited. The role of season of treatment in the efficacy of smoke as a promotive germination agent was investigated over two growing seasons using natural soil stored seedbanks in Banksia woodland near Perth, Western Australia. Smoke was applied to unburnt sites in the autumn, winter and spring of 1994. Germinant emergence and seedling survival of 37 species representing 18 families was monitored in both unburnt sites and in adjacent, recently burnt sites until the second spring after treatment (October 1995). Recruitment from seed was found to be profoundly affected by the season in which dormancy breaking treatment had been applied. The promotive effect extended beyond the initial year of application. For the majority of the species investigated, application of smoke to unburnt sites in autumn promoted a significantly greater germination response than treatment in winter or spring. In only three cases (introduced annuals, the Fabaceae and Hibbenia amplexicaulis) did autumn smoke treatment not yield better germination than in summer-burnt counterparts. However, in almost half of the cases examined, proportions of seedlings surviving past their first summer after emergence in burnt areas were consistently greater than those in smoked or untreated sites. Most notably, no seedlings emerging during the spring of the first year of study survived into the following summer. Implications of the results with respect to future seed bank research and management of native vegetation are discussed.     

腊梅的受精作用及胚胎发生

腊梅 (Chimonanthuspraecox)花两性 ,离心皮雌蕊着生在杯状花托上 ,柱头线形 ,干性。花粉经昆虫传播 ,落在柱头上 1d后萌发 ,第 8d从珠孔进入 ,第 1 4d左右完成双受精 ,为珠孔受精。胚乳为核型胚乳 ;初生胚乳核经短暂休眠进行核分裂 ,位于合点端的游离核首先形成细胞 ,并从合点向珠孔端细胞化 ,第 37d胚乳充满整个囊腔。合子经过近 2周的休眠后开始分裂 ,随着胚的发育 ,大部分胚乳降解 ,为胚的发育提供营养。合点端的胚乳细胞则侵入合点珠心组织 ,为胚进一步发育提供营养。其胚胎发生为柳叶菜型。     

腊梅的受精作用及胚胎发生

腊梅(Chimonanthus praecox)花两性,离心皮雌蕊着生在杯状花托上,柱头线形,干性。花粉经昆虫传播,落在柱头上1 d后萌发,第8d从珠孔进入,第14d左右完成双受精,为珠孔受精。胚乳为核型胚乳;初生胚乳核经短暂休眠进行核分裂,位于合点端的游离核首先形成细胞,并从合点向珠孔端细胞化,第37d胚乳充满整个囊腔。合子经过近2周的休眠后开始分裂,随着胚的发育,大部分胚乳降解,为胚的发育提供营养。合点端的胚乳细胞则侵入合点珠心组织,为胚进一步发育提供营养。其胚胎发生为柳叶菜型。     

EMBRYOLOGY AND DEVELOPMENT OF THE ENDOSPERM HAUSTORIUM OF ARCEUTHOBIUM DOUGLASII

Arceuthobium douglasii develops a dome-like structure, the ovarian papilla, in which 2 megasporocytes are formed. The papilla is not a true ovule, for no integuments are formed, and it is forced aside by the developing endosperm. Megasporocytes are differentiated in the spring, but meiosis does not occur until the following spring. A tetrasporic embryo sac is developed which is 8-nucleate at maturity. Pollination and fertilization occur approximately 13–14 months after initiation of the inflorescence. Only 1 of the 2 embryos develops after fertilization. After fertilization, the embryo sac segregates into 2 parts, one containing the zygote and the disintegrating synergids, the other the primary endosperm nucleus and the degenerating antipodals. This primary endosperm cell elongates toward the base of the ovarian papilla. Cytokinesis then forms an endosperm cell, adjacent to the zygote, and a haustorial cell. The haustorial cell forms several tiers of cells which persist during the development of the embryo and endosperm. The zygote, while still contained within the ovarian papilla, divides, forming a 2-celled sphere. It remains unchanged until after it is conveyed out of the ovarian papilla by the developing endosperm. The development of the embryo and endosperm is arrested in the autumn approximately 3 months after their initiation. They complete their development the following spring and summer.     

Variation between Emex australis populations in seed dormancy/non-dormancy cycles

Abstract Seeds of four Western Australian accessions of Emex australis were stored outside and their germinability tested at 4–6 week intervals for 22 months. Accessions showed cyclical behaviour in germinability, with peaks in autumn/early winter and troughs in spring/summer. There was considerable variation between accessions in dormancy/non-dormancy cycles, but when plants from accessions with contrasting cycles were grown in a common environment, the cycles shown by their seeds were virtually identical. Thus, the parameters of seed dormancy/non-dormancy cycles in E. australis appear to be under environmental control. Only part of an E. australis seed population demonstrated cyclical changes in dormancy status, in contrast to many other annual species whose entire seed populations undergo such changes. Populations also contained individuals which were either continuously dormant or continuously non-dormant following a period of after-ripening. The latter seeds give E. australis the flexibility to recruit opportunistically after summer rainfall events in mediterranean-climate environments.     

Ecophysiology of embryo development and seed germination of the European woodland herbaceous perennial Corydalis cava (L.) Schweigg. & Körte subsp. cava (Fumariaceae)

In this study we examined the germination ecology with special reference to the temperature requirements for embryo development and germination of Corydalis cava subsp. cava, under both outdoor and laboratory conditions. Corydalis cava is a spring flowering woodland tuberous geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, was investigated in a population growing in northern Italy. Immediately after harvest, seeds of C. cava were sown both in the laboratory under simulated seasonal temperatures and naturally. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions, culminating in radicle emergence in winter, when temperatures fell to ca 5°C. Cotyledon emergence also occurred at ca 5°C, but first emergence was delayed until late winter and early spring. Laboratory experiments showed that high (summer) followed by medium (autumn) and low temperatures (winter) are needed for physiological dormancy loss, embryo development and germination respectively. Unlike seeds of C. cava that germinated in winter, in other Corydalis species radicle emergence occurred in autumn (C. flavula) or did not depend on a period of high summer temperature to break dormancy (C. solida). Our results suggest that subtle differences in dormancy and germination behavior between Corydalis species could be related to differences in their geographical distribution.     

A mutational approach to the study of seed development in maize

The maize seed comprises two major compartments, the embryo and the endosperm, both originating from the double fertilization event. The embryogenetic process allows the formation of a well-differentiated embryonic axis, surrounded by a single massive cotyledon, the scutellum. The mature endosperm constitutes the bulk of the seed and comprises specific regions containing reserve proteins, complex carbohydrates, and oils. To gain more insight into molecular events that underlie seed development, three monogenic mutants were characterized, referred to as emp (empty pericarp) on the basis of their extreme endosperm reduction, first recognizable at about 12 d after pollination. Their histological analysis reveals a partial development of the endosperm domains as well as loss of adhesion between pedicel tissues and the basal transfer layer. In the endosperm, programmed cell death (PCD) is delayed. The embryo appears retarded in its growth, but not impaired in its morphogenesis. The mutants can be rescued by culturing immature embryos, even though the seedlings appear retarded in their growth. The analysis of seeds with discordant embryo-endosperm phenotype (mutant embryo, normal endosperm and vice-versa), obtained using B-A translocations, suggests that emp expression in the embryo is necessary, but not sufficient, for proper seed development. In all three mutants the picture emerging is one of a general delay in processes related to growth, as a result of a mutation affecting endosperm development as a primary event.     

羊草受精作用及其胚与胚乳早期发育的观察

利用常规石蜡制片方法研究了羊草受精过程及胚与胚乳的早期发育,其主要结果为：(1)授粉后1h,花粉管破坏1助细胞,释放2精子。精子为眼眉状,难以区分其细胞质鞘;(2)授粉后1～2h,2个精子分别移向卵细胞与极核;(3)授粉后2～3h,精核分别贴附于卵细胞与极核核膜上;(4)授粉后3～10h,精核与卵核融合,并出现雄性核仁,形成合子;(5)授粉后3～4h,精核与极核融合,并出现雄性核仁,形成初生胚乳核,精核与极核的融合比与卵核融合快;(6)传粉后20h,合子分裂,合子的休眠期为10h左右;(7)传粉4h,初生胚乳核分裂,初生胚乳核没有休眠期;(8)羊草双受精作用属于有丝分裂前配子融合类型;(9)胚胎发育属于紫菀型,胚乳发育属于核型胚乳。     

Cytological study of pollen tube growth and early seed development inPetunia inflata

Pollen tube growth from the stigma into the ovule, and the early fruit and seed development following fertilization were examined using fluorescence microscopy, scanning electron microscopy and light microscopy inPetunia inflata. After growing intercellularly in the transmitting tract for 24–36 hr, the pollen tubes emerged into the top part of the ovary cavity and grew along the surface of the septum to reach the ovule. It grew around the furnicle and penetrated the micropyle to enter the embryo sac for fertilization. After fertilization, the endosperm nucleus divided first before the embryo, and the cell wall formation occurred following the division, exhibiting the pattern of cellular type of endosperm development. The first division of the zygote did not occur until 3 days after pollination. At 6 days after pollination, the seeds grew considerably and the endosperm has gone through multiple rounds of cell division. High starch formation in the integument, especially around the embryo sac, was also observed.     

Seed production and outbreaks of non-cyclic rodent populations in deciduous forests

Summary In a 10-year study period, outbreaks of the bank vole, Clethrionomys glareolus, Schreber only occurred in years following huge seed production of European beech, Fagus sylvatica. Intensive winter reprodution preceded the outbreaks, in contrast to a normal breeding season from April through September. No winter reproduction occurred in nearby populations from habitats without mast production. During the winter, the average weight of C. glareolus remained high in the mast forests and the age structure resembled that of a summer breeding population. Despite excess energy requirements of winter breeding, survival rates were similar to that of non-breeding winter populations. In mast years, rodent consumption in the beech forest was estimated as 1.0–10.3% of endosperm production available to postdispersal seed predators. Between mast years rodent consumption made up 30–100% of endosperm production available. Mast years occurred at irregular intervals and seed production seems to be synchronized between individual trees over large areas and induced by climatic events. These phenomena lead to seed predator satiation.     

Double fertilization in Gnetum gnemon (Gnetaceae): its bearing on the evolution of sexual reproduction within the Gnetales and the anthophyte clade

The fertilization process in Gnetum is critical to our understanding of the evolution of sexual reproduction within the Gnetales, a monophyletic group of nonfiowering seed plants that are the closest living relatives to flowering plants. Although much is known about the fertilization process in Ephedra, which is basal within the Gnetales, little is known about sexual reproduction in the derived sister groups Gnetum and Welwitschia. Ovules of Gnetum gnemon were collected at various stages after hand pollination and processed for light, fluorescence, and electron microscopy. Approximately 5 d after pollination, pollen tubes reach sexually mature female gametophytes, which are coenocytic. At that time, a binucleate sperm cell is found within each pollen tube. Within 7 d of pollination, double fertilization events occur when each of two sperm nuclei released from a pollen tube fuses with a separate, undifferentiated female nucleus within the free nuclear female gametophyte, which lacks differentiated egg cells. The products of double fertilization are two viable zygotes; endosperm is not formed. The lack of differentiated egg cells in Gnetum gnemon is unparalleled among land plants and the documentation of a regularly occurring process of double fertilization is congruent with the hypothesis that a rudimentary process of double fertilization evolved in a common ancestor of angiosperms and Gnetales.     

Radicle emergence with increased temperatures following summer dispersal in Trillium camschatcense: A species with deep simple double morphophysiological dormancy in seeds

Seeds with deep simple double morphophysiological dormancy (MPD) need cold stratification during the first winter after dispersal for radicle emergence, followed by the summer for root and bud development and finally a second winter for shoot emergence. In a previous study, we demonstrated that Trillium camschatcense seeds have this type of dormancy with radicles emerging from most seeds after the first winter. However, radicles also emerged from a few seeds in autumn during the same year as dispersal. We thought that temperatures after seed dispersal played a role in radicle emergence before the first winter. To confirm our idea, we investigated germination phenology outdoors, relationships between temperatures after seed dispersal and radicle emergence in the first year outdoors, radicle emergence in the first winter under varied temperatures using incubators, and shoot emergence from seeds with an emerged radicle in the first year outdoors. Our phenology study confirmed that T. camschatcense seeds have deep simple double MPD. Over 7 years, 0.2–7.5% of radicles emerged in the first year before winter and these percentages were moderately positively correlated with temperatures, especially minimum temperatures. Increasing August and September temperatures increased radicle emergence in the laboratory. Shoots emerged from seeds with an emerged radicle in the first year after the first winter. With increased autumn temperatures in warmer regions or with global warming, we predict that germination phenology may shift: increased radicle emergence in the first year and shoot emergence following the first (and not second) winter.     

Observations on Fertilization in Sugar Beet

The whole process of double fertilization in sugar beet has been observed, the main results are as follows: About 2 hours after pollination, the pollen grains germinate, the sperms in the pollen tube are long-oval. 15 hours after pollination, the pollen tube destroys a synergid and releases two sperms on one side or at the chalazal end of the egg cell. The sperms are spherical each having a cytoplasmic sheath. 17 hours after pollination, one sperm enters the egg cell, and the sperm nucleus fuses with the egg nucleus rapidly. 21 hours after pollination, the zygote is formed. In the meantime, the primary endosperm nucleus has divided into two free endosperm nuclei. 25 hours after pollination, the zygote begins to divide, forming a two-celled proembryo. The dormancy stage of the zygote is about 4 hours. In the meantime the endosperm is at the stage of four free nuclei. 17 hours after pollination, the sperm nucleus comes into contact and fuses with the secondary nucleus. The sperm nucleus fuses with the secondary nucleus, faster than the sperm with the egg. he first division of the primary endosperm nucleus is earlier than that of the zygote, it takes place about 20 hours after pollination, the dormancy stage of the primary endosperm is about 2 hours. The endosperm is free nuclear. The fertilization of sugar beet belongs to premitotic type of syngamy. From the stage of zygote to the two-celled proembryo, it can be seen that addition- al sperms enter the embryo sac, but polyspermy has not been observed yet.     

Temperature controls seed germination and dormancy in the European woodland herbaceous perennial Erythronium dens-canis (Liliaceae)

We examined the germination ecology and the temperature requirements for germination of Erythronium dens-canis, under both outdoor and laboratory conditions. E. dens-canis is a spring flowering woodland geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, were investigated in a natural population growing in Northern Italy. Immediately after harvest, seeds of E. dens-canis were either sown on agar in the laboratory under simulated seasonal temperatures or placed in nylon mesh sachets and buried in the wild. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions in the laboratory and in the wild, culminating in radicle emergence in winter when temperatures fell to ≈ 5 °C. Emergence of cotyledons did not occur immediately after radicle emergence, but was delayed until the end of winter. Laboratory experiments showed that temperature is the main factor controlling dormancy and germination, with seeds becoming non-dormant only when given warmth, followed by cold stratification. Unlike seeds of E. dens-canis that germinate in winter, in other Erythronium species radicle emergence occurs in autumn, while in some it is delayed until seeds are transferred from winter to spring conditions. Our results suggest that there is genetic and environmental control of the expression of seed dormancy amongst Erythronium species, which is related to local climate.     

Variation within species and inter-species comparison of seed dormancy and germination of four annual Lamium species

In an ecological context, knowledge of intra-species variation in dormancy and germination is necessary both for practical and theoretical reasons. We used four or five seed batches (replicates) of four closely related annuals co-occurring in arable fields in Sweden: Lamium amplexicaule, L. confertum, L. hybridum and L. purpureum. Seeds used for experiments stemmed from plants cultivated on two sites, each site harbouring one population of each species, thereby ensuring similar environmental history of seeds. Seeds were tested for germination when fresh and after three different pre-treatments (cold or warm stratification, or dry storage) for up to 24 weeks. Seeds were also sown outdoors. Despite substantial intra-species variation, there were clear differences between species. The general seed dormancy pattern, i.e. which environmental circumstances that affect dormancy, was similar for all species; dormancy reduction occurred during warm stratification or dry storage. Even though the response to warm stratification indicates a winter annual pattern, successful plants in Sweden were mostly spring emerged. Germination in autumn occurred, but plants survived winters poorly. Consequently, as cold stratification did not reduce dormancy, strong dormancy in combination with dormancy reduction during dry periods might explain spring germination. It is hypothesised that local adaptations occur through changes mainly in dormancy strength, i.e. how much effort is needed to reduce dormancy. Strong dormancy restricts the part of each seed batch that germinate during autumn, and thus reduces the risk of winter mortality, in Sweden.     

Soil temperatures after the passage of a fire: Do they influence the germination of buried seeds?

Abstract Soil temperatures down to a depth of 5 cm were measured in the days following one fire in summer, one fire in winter and in unburnt vegetation during summer. Soil temperatures did not rise above 40°C after the winter fire or in unburnt vegetation during summer. Consequently, no impact on seed dormancy in the soil seedbank was expected. After a summer fire, soil temperatures above 40°C were found up to 4.5 cm in depth, while temperatures above 60°C were found only in the top 0.5 cm of soil. These temperatures are sufficient to break seed dormancy in some legume species in the seedbank. Hence, the season of burn may influence the number of seeds in the soil that have their dormancy broken and subsequent germination levels.     

Ecological distribution and phenology of an invasive species, Cardamine hirsuta L., and its native counterpart, Cardamine flexuosa With., in central Japan

Cardamine hirsuta is a European species that was recently introduced into Japan and its wide distribution has been confirmed in the Kanto district. To understand mechanisms of the recent spread of C. hirsuta in Japan, a comparative study of the alien species and its native congeneric species, C. flexuosa, was conducted. Habitat preferences, phenology and seed germination were examined. Cardamine hirsuta and C. flexuosa showed distinctive habitat-preferences; the former was most common in open habitats created by recent man-made constructions, and the latter was common in rice paddy fields and surrounding areas. The results indicate that C. flexuosa is a year-long annual, with a mixed phenology of summer and winter germination and growth. Seed dormancy during summer was relatively weak for C. flexuosa, and some plants that germinated early in summer reproduced during the same summer–autumn period. Plants that germinated in late summer and autumn behaved as winter annuals. In rice paddy fields, C. flexuosa is a winter annual because germination is prevented by submergence during summer. Plants flower during the following spring and complete their life cycle before the fields are flooded for rice cultivation. Cardamine hirsuta showed strong seed dormancy during summer and behaved as a typical winter annual. Seeds of C. hirsuta were intolerant to submergence in water, a condition that breaks seed dormancy of C. flexuosa. The results explain the absence of C. hirsuta from rice paddy fields. It was concluded that the spread of C. hirsuta is attributable to the recent expansion of urban habitats created by human activity and has occurred without direct competition with C. flexuosa. Considering recent urbanization in many areas, it is suggested that C. hirsuta has been spreading rapidly in Japan.     

Seasonal differences in mineral content, distribution and leakage of sweet pepper seeds

Pepper seed quality is sensitive to variations in climatic conditions during seed development, which might be associated with accumulation, distribution and leakage of mineral elements from the seeds. This was examined in hybrid seeds of sweet pepper (Capsicum annuum L. cv.‘Hazera’ 1195) in two experiments during two growing seasons. The mean daily temperature (day/night) and daily total radiation receipt during seed development were 27.9/23.2°C and 8.63 kW m^?2 in the summer and 18.3/ 14.9°C and 3.18 kW m^?2 in the winter, respectively. Seeds developed in the summer had lower percentage of seedling emergence and leaked a larger portion (45%) of their K content into the water medium than in winter‐developed seeds. Summer seeds accumulated more K and Cl, but less P, Mg, Ca and the weight ratio of linoleic acid to oleic acid was lower than in the winter seeds. The season did not significantly affect N, S and total fatty acids. The most abundant element on the seed coat surface was K in the summer and Ca in the winter seeds. The cotyledon and endosperm of the summer seeds contained relatively higher ratios of K and Ca and lower ratios of P and Mg than the winter seeds. Transportation of mineral nutrients appeared to be involved in the effect of heat and moisture stresses on emergence quality of the pepper seeds.     

Seasonal cycle of seed dormancy controls the recruitment of Butia odorata (ARECACEAE) seedlings in savanna‐like palm tree formations in southern Brazil

Butia odorata (Barb. Rodr.) Noblick is a palm tree that grows in savanna‐like formations in subtropical regions of South America, and whose regeneration is threatened by agricultural management. Its diaspores are dormant after dispersal which takes place during the summer and early autumn. The aim of this study was to investigate seasonal and microhabitat effects on the germination and seedling recruitment of this palm species. Diaspores were sown in the field, in both open lands and forest patches. During 2 years, we measured seed germination, viability and moisture, seedling emergence and germination response to warm stratification of those seeds that failed to germinate in the field. Germination was concentrated during the summer, when soil temperatures were highest, whilst seedling emergence peaked in the autumn and early winter, when temperature and humidity conditions became less extreme. In open lands, there were two pulses of germination (first and second summer), whilst in forest patches, a single pulse (second summer) was detected. Although overall germination did not differ between microhabitats, the percentage of seedling emergence from seeds that remained buried until the end of the experiment was almost twice as large in the forest patches compared with open areas. The viability of seeds declined over time, particularly in open areas. Laboratory‐induced warm stratification was found to act on seed dormancy release in a cyclic way, being far more effective on seeds retrieved from the field in spring–summer months than in those retrieved in the winter. This cyclic pattern of dormancy in B. odorata seeds results in major seedling recruitment after the summer, under wetter and cooler conditions, thus reducing mortality risk. This process can be enhanced by the presence of surrounding vegetation, which both increases seedling emergence and/or prolongs seed viability.     

Changes in dormancy levels ofFraxinus excelsior L. embryos at different stages of morphological and physiological maturity

The dormancy status ofFraxinus excelsior embryos at different developmental stages under environmental conditions was examined over a period of 2 years. For each sampling date the length of the fruit, of the seed, and of the embryo were measured, and the embryological stage determined. The depth of dormancy was assayed by the germination behaviour of isolated embryos under aseptic conditions on an agar medium without nutrients. As an approach towards a quantitative estimate of the dormancy status, the degree of inhibiton of germinative growth in the embryonic organs was evaluated on the basis of four categories from none to full germinative growth. From these ratings a dormancy index was calculated, expressing the mean dormancy status of the embryos at a given date. Embryo dormancy already became apparent during embryogenesis and reached its highest level during the later phase of reserve deposition in the seed. A marked loss of embryo dormancy occurred during the phase of maturation drying in autumn, followed by a moderate increase in winter. In hydrated seeds in spring the embryo was gradually released from dormancy and enlarged further. In maintaining the embryo ofF. excelsior in a developmental but not germinative mode, dormancy mechanisms within the embryo and the endosperm, combined with environmental factors, may be involved.