CAM induction by photoperiodism in green callus cultures from a CAM plant

Abstract Green calli obtained from leaves of the CAM-inducible plant Kalanchoe blossfeldiana cv. Montezuma were grown either under long-day or short-day regimes in the Phytotron of Gif-sur-Yvette. Callus cells were found to be CAM inducible by the short-day treatment, according to levels and diurnal oscillations of malate pools and phosphor-enolpyruvate (PEP) carboxylase (EC.4.1.1.31) capacity. De novo synthesis of PEP carboxylase was shown to occur under the short-day regime. In spite of continuous net CO² output, CAM-like patterns of CO₂ exchange by calli were obtained under the short-day treatment. After 2 months under both photoperiodic conditions, spontaneous organogenesis in callus tissues gave rise to numerous shootlets which were determined as photoperiod dependent for CAM; the same was so for plants originating from these shootlets.     

Photoperiodic regulation of receptacle induction in Sargassum horneri (Phaeophyceae) using clonal thalli

We developed a clonal culture of Sargassum horneri to investigate the effect of photoperiod on reproduction in this species. Regenerated vegetative thalli were obtained using lateral branches excised from a thallus grown from a single embryo under short‐day conditions (SD = 10:14 h light : dark cycle). Lateral branches excised from the SD‐regenerated thallus became vegetative thalli that remained in that phase as long as they were cultured under SD. When an excised lateral branch was cultured under long‐day conditions (LD = 14:10 h light : dark cycle), it began to enter the reproductive phase while still less than 50 mm long. Induction of the reproductive phase was accompanied by a distinctive morphological change – suppression of blade formation at the apical region of the branch; elongation of branches without blades was then followed by differentiation of receptacles bearing conceptacles on their surface. Apices of receptacles were able to interconvert between reproductive and vegetative phases, as blades resprouted upon transfer from LD to SD. The critical day length for induction of receptacle formation was between 13 and 14 h; receptacle formation was also induced under SD conditions with night breaks (NBs). These results strongly suggest that reproductive regulation of S. horneri is a photoperiodic long‐day response. NBs with blue and green light were effective for reproductive induction but not with red light. This suggests that blue‐ and/or green‐light photoreceptors are involved in the photoperiodic reproductive response of S. horneri.     

Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.     

Effects of photoperiod and temperature on blood feeding,oögenesis and fat body development in the mosquito, Culiseta inornata

In the laboratory, blood feeding rates, oögenesis and fat body development were utilized to assess the effects of photoperiod and temperature in the conditioning of adult female Culiseta inornata for aestivation. Long-day (16L:8D) and short-day (8L:16D) photoperiods were examined in combination with temperatures of 15°C, 20°C, and 25°C. All tested individuals were reared from egg to adult under one of six possible regimes.Blood feeding occurred in approximately 60% of the females subjected to short-day conditions. Females subjected to long-day regimes exhibited gonotrophic concordance (the inhibition of blood feeding), evidenced by a decrease in overall blood-feeding rate to 20%. Inhibition of blood feeding by long-day females decreased in a linear fashion with increasing temperature. No interaction of the effects of photoperiod and temperature upon blood feeding was apparent. Examination of the ovarioles of post-blood feeding females, reared under each of the above conditions, revealed no evidence of gonotrophic dissociation (the inhibition of oögenesis despite continued blood feeding).Fat body hypertrophy occurred in females reared under long-day conditions, whereas hypotrophy of this tissue was apparent in short-day females. No significant difference in fat body development occurred between parous and nulliparous females reared under long-day photoperiod conditions. Short-day blood-fed females reared at 15°C deposited a significantly greater amount of fat than short-day blood fed females reared at 20° and 25°C, and short-day nonbloodfed females reared at 15°, 20°, and 25°C. The primary stimulus for fat body hypertrophy appears to be long-day photoperiod conditions. Temperature exerted little discernible effect upon this process, and there was no interaction between the effects of photoperiod and temperature upon the degree of fat body development.     

Life history of Japanese Stypocaulon durum (Sphacelariales, Phaeophyceae)

Phenology, morphology, life history and responses to different temperature and photoperiod conditions were studied in Japanese Stypocaulon durum (Ruprecht) Okamura. Erect thalli of the species were collected year-round, but the mature thalli forming either uniloc-ular sporangia or two different types of plurilocular structures (evidently gametangia) on separate thalli were found only in winter. ln culture, an isomorphic life history is suggested for the species, alternating between a sporophyte forming unilocular sporangia and gam-etophytes forming plurilocular macro- (female) and micro- (male) gametangia. Contents of unilocular sporangia were not released, but germinated in situ, developing into erect thalli forming plurilocular gametangia. Macrogametangia released aplanogametes (oospores), but male gametangia appeared to be non-functional, although flagellated cells were once formed in the loc-uli. This is the first report of plurilocular gametangia in the species. Although the species grew well and matured under considerably lower temperature conditions than European Stypocaulon scoparium (L.) Sauvageau, its temperature requirements showed similarity to northwestern Atlantic Stypocaulon species. This supports the notion that northwestern Atlantic Stypocaulon is conspecific with S. durum.     

The effect of artificial photoperiod on growth and reproduction in the land snail Cepaea nemoralis

Summary

Specimens of Cepaea nemoralis were raised from egg to adult in the laboratory for 15.5 months in two artificial photoperiods: short-day (LD 8:16) and long-day (LD 16:8). Over 20% of the snails in each photoperiod were functional adults by the end of the experiment and had fully developed male and female reproductive systems as determined both by dissection and by oviposition. A total of 925 eggs were laid by animals reared in a long-day photoperiod, whereas animals in a short-day photoperiod laid 677 eggs. Animals reared in a long-day photoperiod grew slightly faster (2.04 mg/day compared to 1.73 mg/day), began laying eggs about 18 days sooner, and suffered higher mortality (19% compared to 11%) than animals reared in a short-day photoperiod. These results clearly establish for C. nemoralis that gametogenesis and the development of a functional hermaphroditic reproductive system are neither prevented nor significantly retarded by exposing the animals to a short-day photoperiod for their entire pre-reproductive life.     

The life history of Ahnfeltiopsis paradoxa (Gigartinales, Rhodophyta) in laboratory culture

The life history of the red alga Ahnfeltiopsis paradoxa (Suringar) Masuda (Phyllophoraceae, Gigartinales) from Japan was completed in laboratory culture. Carpospores isolated from field-collected plants germinated to form circular crusts that were composed of a monostromatic hypothallium consisting of radiating filaments, a polystromatic perithallium consisting of tightly coalescent erect filaments, and hypobasal tissue derived from the hypothallium. The crusts were induced to sporulate by transferring them from short-day to long-day regimes at 15° and 2°C. Each crust produced several nemathecia along 1-4 concentric rings. Intercalary, cruciately or decussately divided tetrasporangia were formed in 4-6 (1-2 at the margin of the nemathecium) successive cells of a single filament of the nemathecia. Tetraspore germlings gave rise to basal discs from which upright axes developed. The upright axes first grew without branches or were sparsely branched and later bore many marginal reproductive proliferations. Procarps and spermatangia were formed in the proliferations on different individuals. Carposporophytes developed on female plants that were co-cultured with male plants. Gonimoblast filaments were formed from an auxiliary cell that fused with a carpogonium. Carposporangia developed from gonimoblast filaments and medullary cells contacted by the gonimoblast filaments. Carpospores were discharged through carpostomes formed in the thickened cortex. Tetraspores were cultured from field-collected crusts of a morphology similar to that of cultured tetrasporophytes. They gave rise to upright gametophytic axes similar in morphology to this species as seen in the field.     

LIFE HISTORY OF COLPOMENIA SINUOSA (SYCTOSIPHONACEAE,PHAEOPHYCEAE) IN THE AZORES1

Colpomenia sinuosa (Mertens ex Roth) Derbès and Solier (Scytosiphonaceae, Phaeophyceae) is a common species on the rocky intertidal shores of the Azores, where reproductive gametophytes occur throughout the year. Life‐history studies of this species were carried out in culture, and both sexual and asexual reproduction were observed. Anisogamous gametes fused to form zygotes. The zygotes gave rise to a filamentous prostrate sporophyte generation bearing unilocular sporangia, under both short‐day and long‐day conditions at 15 and 22°C, and to both unilocular and plurilocular sporangia, under the lower temperature condition. Unispores developed into gametophytes, and plurispores gave rise to filamentous sporophytes. Asexual reproduction was carried out by unfused female gametes and asexual plurispores produced from the same gametophyte. Unfused gametes developed into filamentous prostrate sporophytes producing unilocular sporangia in both culture conditions, and unispores released from the sporangia gave rise to gametophytes. Asexual plurispores from field gametophytes, under both culture conditions, developed directly into new gametophytes. The species exhibited three types of life history: a heteromorphic, diplohaplontic; a heteromorphic, monophasic (both with alternation between the erect and filamentous prostrate thalli); and a monomorphic, monophasic.     

CIRCANNUAL GROWTH RHYTHM AND PHOTOPERIODIC SORUS INDUCTION IN THE KELP LAMINARIA SETCHELLII (PHAEOPHYTA)1

Growth and reproduction of laboratory-grown sporophytes of Laminaria setchellii Silva were investigated in a tank system with controlled conditions of daylength, temperature, and nutrients (N and P). A circannual growth rhythm of the frond was detected under constant laboratory conditions. In continuous long-day and night-break conditions the period τ of the free-running rhythm varied between 11.3 and 17.3 months; in short-day conditions the frond grew indefinitely. The growth rhythm of individual plants could be synchronized by a simulated annual cycle of day-length with a period of T = 12 months. The four seasons of the year were simultaneously simulated by phase shifting the annual cycle of daylength by 3, 6, or 9 months in three out of four tanks. The annual growth cycle followed these phase shifts, and initiation of the new blade always started just after the winter daylength minimum. The formation of sori was induced by a genuine photoperiodic short-day reaction in 1- to 2-year-old plants. Sori became, visible 9–14 weeks after transfer of individual plants from long-day to short-day conditions, whereas plants cultured in continuous long-day or night-break conditions remained sterile. Sporophytes with or without blades were able to continue growth or produce new blades in continuous darkness.     

Effects of Root Excision on Swelling of Leaf Sheath Cells and on the Arrangement of Cortical Microtubules in Onion Seedlings

Unlike abundantly-leafed mature plants, 2- to 3-leafed youngonion seedlings (Allium cepa L. cv. Senshu-Chuko) showed noswelling of leaf sheaths even when they were kept under long-dayconditions. When their roots were excised, however, the seedlingsshowed sign of swelling. Before swelling became evident, changesin the arrangement of cortical microtubules occurred in leafsheath cells. The microtubules, which were oriented transverselyto the cell axis in unexcised seedlings, were oriented longitudinallyor obliquely in excised seedlings. Such effects of root excisionwere observed only in seedlings grown under long-day conditions,but not under short-day conditions. (Received July 14, 1984; Accepted September 26, 1984)     

Short-day and low-temperature conditions promote reproductive maturation in the terrestrial slug, Lehmannia valentiana

Terrestrial slugs (Lehmannia valentiana) collected from a field in Osaka, southwestern Japan, were reared under long-day (16-h light and 8-h darkness, LD 16:8) or short-day conditions (LD 12:12) at 15, 20, or 25 degrees C for 60 days. Slugs reared under short-day conditions were heavier than those reared under long-day conditions; however, slugs reared at 15 degrees C gained more weight regardless of the photoperiodic condition. Gonads were significantly heavier under short-day conditions than under long-day conditions, and oogenesis and spermatogenesis were also induced under short-day conditions. Under short-day conditions, reproductive maturation was suppressed at 25 degrees C as compared with 15 and 20 degrees C, whereas under long-day conditions, lower temperatures induced reproductive maturation. In contrast, slugs reared under short-day conditions at 20 degrees C from the time of hatching gained more weight than those reared under long-day conditions at the same temperature. Moreover, short-day conditions induced reproductive maturation, similar to that observed in the field-collected slugs. In conclusion, short-day and low-temperature conditions promoted growth and reproductive maturation, whereas long-day and high-temperature conditions suppressed them in L. valentiana. Thus, L. valentiana reproduces from late autumn to spring in Osaka.     

Relationship between photoperiod and secretion of the diapause hormone during larval stages of the silkworm, Bombyx mori L., reared on an artificial diet

The suboesophageal ganglion of the silkworm, Bombyx mori synthesizes sufficient diapause hormone to produce diapause eggs, regardless of the photoperiodic conditions experienced during the larval stages. When larvae destined to produce non-diapause eggs are implanted with the brain-suboesophageal ganglion complex from larvae which have been reared under short-day conditions, the resulting adults lay diapause eggs. The larvae receiving the complex from larvae reared under long-day conditions gave rise to adults which did not produce any diapause eggs. The brains from pupae which have been reared under long-day conditions show an activity inhibiting the secretion of diapause hormone by the suboesophageal ganglion. The mechanism through which the brain controls the secretion of diapause hormone from the suboesophageal ganglion can be modified by photoperiodic conditions during the larval stages.     

LIFE HISTORY AND SYSTEMATIC POSITION OF PSEUDOCHORDA GRACILIS SP. NOV. (LAMINARIALES,PHAEOPHYCEAE)1

Pseudochorda gracilis sp. Nov. (Pseudochordacease, Laminariales) is described from the Japan Sea coast of Hokkaido/ the species is subtidal, epilithic and annual, appearing in spring and maturing in winter. Erect thalli grow solitary or in tufts on a small discoid holdfast. They are simple, cord-shaped and hollow, with inner hyphal filaments, cylindrical medullary cells and paraphyses consisting of 3–6 cells. Hair tufts are observed only in young thalli. Unilocular sporangia are sessile and narrowly ovate. In culture, P. gracilis shows a heteromorphic life history with oogamy, characteristic of the order Laminariales. Gametophytes are dioecious and dimorphic. Gametophytes mature under lower temperature conditions (usually below 10°C), and sporophytes mature under low temperature and short-day conditions (5°C, SD). The seasonal growth pattern of the species results from the photoperiod-temperature conditions controlling saprophyte maturation.     

Control of diapause induction by a change in photoperiod in Melanoplus sanguinipes

The effect of photoperiod on diapause induction, developmental time and body size was examined in Melanoplus sanguinipes, the lesser migratory grasshopper. Contrary to what is found in most insects, facultative diapause-egg production is found to be controlled by changing rather than constant photoperiods. In addition, developmental time is shown to be faster under short-day photoperiods than long-day photoperiods. And finally, body size is larger under long-day photoperiods and smaller under short-day photoperiods. Implications of these results for the regulation of the seasonal life cycle of M. sanguinipes in the field are discussed.     

Alternative life cycles controlled by temperature and photoperiod in the oligophagous bug, Dybowskyia reticulata

Abstract. Effects of temperature and photoperiod on the induction and re-induction of adult diapause were examined in Dybowskyia reticulata (Dallas) (Heteroptera: Pentatomidae). Adults collected from the field after overwintering in early summer continued oviposition under long-day conditions of LD 16:8 h at 20 or 25°C, while they re-entered diapause under short-day conditions of LD 12:12 h at 25, 27.5 or 30°C. By contrast, adults reared in the laboratory from eggs at 20 or 25°C entered diapause under both long-day and short-day conditions, whereas those reared at 27.5 and 30°C entered diapause only under short-day conditions. Under quasi-natural conditions in 1993, when summer temperature was low, most adults of the first generation entered diapause in late July. However, in the warmer summer of 1996, oviposition was recorded in many females that ecdysed into adults from July to early August. Even though the seeds of the host plants occur in a restricted period from early summer to early autumn, in warmer years D. reticulata may produce a second generation. The response to temperature with a threshold between 25 and 27.5°C in D. reticulata brings about a switch between the univoltine and bivoltine life cycles.     

Summer dormancy ensures univoltinism in the predatory bug Picromerus bidens

The seasonal cycle of Picromerus bidens L. (Heteroptera: Pentatomidae) is usually considered to be univoltine with an obligatory winter egg diapause. Seasonal adaptations of the species were studied in the laboratory and in field experiments. When reared under short-day photoperiodic conditions (L12:D12 and L14:D10), all females began to lay eggs synchronously soon after their emergence. However, in the females reared under long-day conditions (L18:D6 and L20:D4) and outdoors in June–July, oviposition was significantly delayed. This delay in reproduction induced by photoperiodic conditions and then spontaneously terminated was considered to be aestivation. Egg batches laid by females in the laboratory and in the field were kept at 25 °C for two months. From 30.8 to 93.8% of batches contained eggs which hatched without cold treatment between day 14 and 60 after oviposition. The proportion of eggs hatched was 17.7 to 20.9% in the short-day regimes, while it was significantly less (5.7 to 6.0%) under long-day conditions. It is concluded that in some eggs diapause is of low intensity and that if under natural conditions the first batches had been laid at the end of June, nymphs would have hatched at least from some eggs during the same season even without cold treatment. Such untimely hatching would have resulted in the death of nymphs and adults unprepared for overwintering. A photoperiodic response which induces aestivation in the early emerging adults in June–August may prevent early oviposition and occurrence of a second generation and thus maintains univoltinism in P. bidens.     

Development Times Under Long- and Short-Day Conditions in the Kanzawa Spider Mite <Emphasis Type="Italic">Tetranychus kanzawai</Emphasis> (Acari: Tetranychidae)

In a theoretical framework, the development time of arthropods is closely associated with population growth, so that strong selection pressure may be imposed on development patterns. In this context, selection towards shorter development times may act on female mites in the season when population size is growing. In contrast, such selective pressure may be weaker just before hibernation, because the stage adjustment may be more important than population growth in such seasons. Thus, selection to shorten development times may act more strongly in long daylength conditions, so that different developmental responses among the seasons (reaction norm) may evolve. I predicted that the development time is shorter under the long-day conditions of population-growing seasons than under the short-day conditions of hibernating seasons, and that genetic variation at the long-day is smaller than that at the short-day. In order to test this prediction, the egg-to-adult development time of a population of Tetranychus kanzawai (Acari: Tetranychidae) was investigated under short (9L:15D) and long (15L:9D) daylength conditions at 18 °C. In addition, genetic variation in female development times under these conditions was estimated by using half-sib design. The development time of females at short day was 1 day longer than at long day under the experimental conditions. Male development times follow the pattern of females with <1 day delay. However, heritabilities under these two treatments were quite low and neither was significant. Thus, only the first prediction was supported by the results. In addition, maternal and/or non-additive effects in the female development time were highly significant under both daylength conditions examined.     

LIFE HISTORY AND GROWTH OF LAURENCIA BRACHYCLADOS (RHODOPHYTA,CERAMIALES)1

Laurencia brachyclados Pilger from Hawai'i completed a “Polysiphonia-type” tri-phasic life history in 21 weeks in laboratory culture. Tetraspores developed into gametophytes in a nearly 1:1:1 ratio of females: males: non-reproductive. Carpospores were released as early as 21 days after mixing virgin female and male gametophytes. Cultured thalli showed a “guerilla type” growth form. Other Hawaiian Laurencia species in culture had longer maturation times or remained non-reproductive. Variation in life history schedules may influence Laurencia species coexistence and algal community structure.     

RESPONSE TO PHOTOPERIOD AND TEMPERATURE BY SPARTINA ALTERNIFLORA (POACEAE) FROM NORTH CAROLINA AND SPARTINA FOLIOSA FROM CALIFORNIA

Seedlings of Spartina alterniflora Loisel. from Oregon Inlet, North Carolina were generally taller and produced significantly more culms and total dry weight than those of S. foliosa Trin. from two California populations (Alameda Beach and Marin County) in eight photoperiod-temperature treatments over a 17-wk period. Seedlings of S. alterniflora produced maximal biomass in 30–26 C whereas those of S. foliosa produced maximal biomass in 22–18 C, both under long-day conditions. The average photosynthesis rate for S. alterniflora (3.0 mg C g dry wt^−- hr^−-1) was 1.6 times higher than that for S. foliosa (1.9 mg C g dry wt^−-1 hr^−-1), but the dark respiration rates (0.3 and 0.4 mg C g dry wt^−-1 hr^−-1, respectively) were not significantly different. The proportion of rhizomes was greater under short than long-day conditions for both species in most temperature regimes. The average shoot soluble carbohydrate and starch concentrations were higher for S. foliosa (8.4 and 0.9%, respectively) than those for S. alterniflora (6.0 and 0.4%, respectively). The average rhizome soluble carbohydrate concentration (18%) for S. alterniflora under long-day conditions was significantly lower than that under short-day conditions (28%) and also lower than those for S. foliosa under both long (26%) and short-day (25%) conditions. Rhizome starch concentrations of S. alterniflora were significantly higher in the short than in most long-day temperatures and were generally higher for S. alterniflora than for S. foliosa under short-day conditions. The root starch concentration of S. alterniflora under short-day conditions (1.3%) was higher than that under long-day conditions (0.2%) and also higher than those of S. foliosa under both long (0.2%) and short-day (0.7%) conditions. The two species exhibited similar patterns of carbohydrate storage in belowground organs, similar flower initiation processes not under strict photoperiod control, and similar respiration rates, but significantly different photosynthetic rates and growth responses with S. alterniflora having the potential to be the more productive species.     

Phytotron experiments in Pisum

Summary The flowering behavior of 59 Pisum mutants and 228 recombinants was studied in the phytotron in four different photoperiods (continuous light, long-day 18/6 h, short-day 12/12 h, extreme short-day 6/18 h). There was no or little difference in the response of the genotypes to long-day and permanent light, whereas great differences were observed between long- and short-day 12/12 h and between the two short-day trials. About half the genotypes tested were unable to survive or to flower in extreme short-day. Some recombinants, however, had an almost normal development under these unfavorable conditions. Gene fis controls the photoperiodic reaction of the plants: they are unable to flower in short-day. Gene fds negatively influences gene efr for earliness: it causes a strong delay of flowering of efr recombinants in long-day and suppresses the formation of functionable flowers in short-day. Most of the genotypes tested showed a specific reaction to the four photoperiods different from that of the mother variety and the other genotypes. The practical aim of our phytotron experiments is the preselection of Pisum genotypes which might be suited for cultivation in countries with short-day climate.This paper is dedicated to Professor Karl-Ernst Wohlfarth-Bottermann on his 65th birthday