THE INITIATION AND ELONGATION OF RHIZOID CLUSTERS IN CAULERPA PROLIFERA

Under improved environmental conditions the average rate of elongation of the rhizoid cluster was 5.2 mm/day in the first 3 days. The rate decreases thereafter and is close to zero around 5–6 days after initiation. The elongation, like that of the rhizome, is chiefly by means of “tip-growth.” The presumptive site of the initiation is 1.3–1.7 mm from the rhizome tip. Under present experimental environment, the presumptive time of the initiation was mainly at 1 to 3 hr after the beginning of the light period. Initiation in the dark period was less frequent, and little or no initiation in the middle and later parts of the light period was observed. The average plastochron of the cluster was 0.91 day. The rate of elongation of the rhizoid cluster was less variable than that of the rhizome. Hence, the slope of calculated regression line from the plot, the cluster length vs. its distance from the rhizome tip, is steeper when the rhizome grows slower (4.9–5.8 mm/day) and less steep when the rhizome grows faster (7.3–8.9 mm/day). In spite of some variation in the slope of the regression line, the intersection between the line and X-axis remains about the same. The application of information on the presumptive site and presumptive time of the rhizoid cluster initiation to a developmental study of micro-events taking place prior to the morphological differentiation is proposed.     

Rhizome gravitropism precedes gravimorphogenesis after inversion of the green algal coenocyte Caulerpa prolifera (Caulerpales)

Previous investigations reported that the marine alga Caulerpa prolifera, a giant coenocyte, had a fast morphogenetic response to a change in position with regard to gravity: the next rhizoid formed approximately 1 day later on the new underside of the inverted rhizome (Jacobs and Olson, 1980, American Journal of Botany 67: 141–146). Preceding the change in site of rhizoid development was a striking accumulation of amyloplasts at the new prospective rhizoid-initiation site in the rhizome tip. Detailed monitoring with video equipment of inverted Caulerpa plants, growing under controlled conditions, now reveals that after the amyloplast accumulation, but before the gravimorphogenetic effect on rhizoid development, the inverted rhizome tip shows a negative gravitropism that restores it to its normally upturned position. Because the experiments were performed with two-sided illumination, this was clearly not a phototropic response.     

DEVELOPMENTAL CHANGES IN THE ALGAL COENOCYTE CAULERPA PROLIFERA (SIPHONALES) AFTER INVERSION WITH RESPECT TO GRAVITY

The algal coenocyte Caulerpa prolifera produces three types of organs, each with a different orientation. The effect of gravity in controlling the development of these organs was investigated. We inverted plants by rotating them 180° around the horizontal rhizome axis, then compared development of the inverted plants with plants matched by size and differentiation. Quantitative data were obtained from photographic records. After inversion the site of organ differentiation was changed with no change in the timing: the next rhizoid (and all subsequent ones) differentiated on the “now-under side” of the inverted rhizome, the next leaf formed on the “now-upper side.” Despite the fast change in the site of organ differentiation, other parameters were not changed by inversion (e.g., rate of elongation of rhizomes or leaves, rate of formation of leaves or rhizoids). Because such changes also occurred in plants with balanced lighting from two sides, it is clear that gravity alone can control these changes without reinforcement from top illumination. After leaves were initiated, they did not change their orientation, showing neither positive phototropism (at our light levels) nor negative geotropism, even when elongating substantially. Torsion of the rhizome tip did not precede the change in site of rhizoid initiation.     

Effects of continuous illumination on the rhizoid cluster of Caulerpa prolifera

Under a photoperiodic regime of 12-hr light and 12-hr dark (12L-12D)and continuous illumination (24L-0D), at 24.0±1°C,the rate of elongation of the rhizoid cluster was 8.1 mm/day,and the presumptive site of the cluster initiation was 1.1–1.9mm from the rhizome tip. However, the plastochron interval under24L-0D regime was shorter (0.87 day) than under 12L-l2D regime(1.18 days). Under both regimes, the plants tended to adjusttheir presumptive time of cluster initiation to be in the twoperiods of the day, 0400–0900 hr and 1800–2100 hr.None of the following seemed to correlate with the initiationof a new cluster: a certain number of hours after the initiationof the youngest cluster, a certain distance on the rhizome distalto the youngest cluster, the youngest cluster reaching to acertain length, or a certain value of the rhizoid plastochronindex. This adjustment by the plants thus suggested the plantshaving an ability to perceive the two preferred periods forcluster initiation. ¹ Report of work supported by research grant GB-7885 from theNational Science Foundation, and in part by Research Councilof Rutgers University. (Received June 2, 1972; )     

ON THE CORRELATION OF PRIMARY ROOT LENGTH,MERISTEM SIZE AND PROTOXYLEM TRACHEARY ELEMENT POSITION IN PEA SEEDLINGS

Pea seedlings (Pisum sativum L. cv. Alaska) were darkgrown in vermiculite. Roots of various lengths were cleared, stained and measured to determine the relative meristem height (MH), width and volume and the distance to the most proximal protoxylem tracheary element (PTE). A correlation was found between root length, MH and PTE position as follows: in roots from 4–40 mm as the root elongated the MH lengthened and PTE position increased its distance from the body/cap juncture; in roots 40–80 mm MH and PTE position remained approximately constant; in longer roots (80–120 mm) MH became shorter, and PTE position closer to the tip as the root elongated. The relationship, using our measurement procedure was that for every 0.19 mm change in MH, the PTE position changed by 1 mm. This response was partially growth rate dependent since short roots (4–80 mm) grew at a constant rate and longer roots (80–140 mm) grew slower. Root manipulations and trifluralin treatment, to inhibit cell division, caused tip swelling and modulated the position of the PTE toward the root tip. The control of the spatial relationship between meristem size and maturation position is discussed.     

Accumulation of amyloplasts on the bottom of normal and inverted rhizome tips of Caulerpa prolifera (Forsskål) Lamouroux

Caulerpa prolifera (Chlorophyta) exhibits a gravimorphogenetic response to inversion by switching the site of new rhizoid initiations to correspond with the new direction of the gravity stimulus. When plants were fixed at 6 and 24 h after being held in either a normal or an inverted position the switch in the site of organ differentiation, upon inversion, was found to be preceded by the accumulation of starch-containing amyloplasts on the bottom of the rhizome. Approximately 1.5–2.0 times as many amyloplasts were found at the bottom of normal or inverted rhizomes as compared with the top in a region extending from 200 to 1,000 μm behind the rhizome tip. All new rhizoid initials were located in the region of amyloplast accumulation and each rhizoid initial contained numerous amyloplasts.     

Meristem culture and virus eradication in Alstroemeria

Stem apical meristems, rhizome apical meristems and rhizome axillary meristems excised from Alstroemeria plants were grown in vitro on modified Murashige and Skoog (MS) media containing different concentrations of gibberellic acid and 6-benzylaminopurine (BA). Plantlets developed from stem apical meristems never regenerated a rhizome and eventually died. The highest regeneration rate (74.1%) of plantlets with a rhizome was observed when rhizome axillary meristems were grown on modified MS medium containing M 8.9 of BA. Alstroemeria mosaic potyvirus (AlMV) could be eradicated from infected Alstroemeria plants through meristem culture. The rate of virus eradication was 73.7 and 14.7% for plantlets developing from explants measuring 0.7 mm and 2.0 mm, respectively. Greenhouse evaluation of virus-negative and AlMV-infected Alstroemeria plants showed that healthy plants produced more floral stems, more vegetative stems, longer floral stems and gave a higher fresh weight than infected plants.     

Membrane recycling occurs during asymmetric tip growth and cell plate formation inFucus distichus zygotes

Summary Zygotes of the brown algaFucus distichus undergo a series of intracellular changes resulting in the establishment of a polar growth axis prior to the first embryonic cell division. In order to examine the dynamics of membrane recycling which occur in the zygote during polar growth of the rhizoid, we probed living Fucus zygotes with the vital stain FM4-64, N-(3-triethylammoniumpropyl)-4-(6-(4-(diethylammo)phenyl)hexatrienyl)pyridinium dibromide. In newly fertilized, spherical zygotes, FM4-64 staining is symmetric and predominantly in the perinuclear region which is rich in endoplasmic reticulum, Golgi, and vacuolar membranes. As rhizoid or tip growth is initiated, this population of stained membranes becomes asymmetrically redistributed, concentrating at the rhizoid tip and extending centrally to the perinuclear region. This asymmetric localization is maintained in the zygote throughout polar growth of the rhizoid and during karyokinesis. Subsequently, FM4-64 staining also begins to accumulate in a central location between the daughter nuclei. As cytokinesis proceeds, this region of stain expands laterally from this central location, perpendicular to the plane of polar rhizoid outgrowth. The staining pattern thus delineates the formation of a cell plate, similar spatially to the accumulation of nascent plate membranes of higher plants. Treatment of Fucus zygotes with brefeldin-A inhibits both asymmetric growth of the rhizoid and formation of a new cell plate. These data suggest that inF. distichus FM4-64 is labeling a Golgi-derived membrane fraction that appears to be recycling between the site of tip growth, perinuclear region, and new cell plate.Abbreviations AF after fertilization - ASW artificial seawater - BFA brefeldin A - ER endoplasmic reticulum - FM4-64 N-(3-triethylam-moniumpropyl)-4-(6-(4-(diethylamino)phenyl)hexatrienyl)pyridinium dibromide     

Taxonomic notes on three species of Herposiphonia (Ceramiales, Rhodophyta) found in Japan

Three species of the red algal genus Herposiphonia (Ceramiales, Rhodomelaceae) found in Japan are described, and taxonomic features of the genus are discussed. Herposiphonia crassa Hollenberg is reported from Japan for the first time and is characterized by thick axes (200–350 µm in diameter) and determinate branches (100–200 µm in diameter), relatively short determinate laterals (400–1200 µm in length) with a large number of periaxial cells (15–19 per segment) and three (occasionally two or four) vigorously developed (1.8–2.5 mm in length by 50–75 µm in diameter basally) trichoblasts on each determinate lateral. Herposiphonia elongata Masuda et Kogame is also reported from Japan for the first time and is characterized by the conspicuous thickening growth of cystocarp‐bearing branches and spermatangial branches with an elongated sterile tip. Some newly found features of Herposiphonia fissidentoides (Holmes) Okamura are presented: the rhizoid production from the central portion of parental periaxial cells in addition to the distal end, virtual absence of vegetative trichoblasts, production of procarpial trichoblasts and spermatangial branches on fertile determinate branches on short indeterminate laterals, cystocarps sometimes with a short spur, and extremely large tetrasporangia.     

DEVELOPMENTAL STUDIES ON THE COENOCYTIC ALGA, CAULERPA SERTULARIOIDES

Thallus growth and development in the coenocyti alga Caulerpa sertularioides (Gmelin) Howe have been studied quantitatively. In unsupplemented seawater at 26 C and in a 12:12 hr light/dark cycle, new rhizomes formed near the old growing points on thalli transplanted from the ocean. Adjacent to the apices of new rhizomes, rhizoids were produced downward, regularly spaced and at a rate of about 1.5/day. Upright. shoots developed irregularly in acropetal succession behind the tips of either the parent or the new rhizomes, which arose as branches of the old. Rates of rhizome, rhizoid, and upright shoot elongation were 0.4, 0.81, and 0.54 cm/day, respectively. Thalli survived up to 2 months in unsupplemented seawater. Long-term growth was obtained by varying culture conditions. A substratum of sand, apparently rich in microorganisms, produced long-term thallus growth in seawater, and the form of development changed so that upright shoot formation was promoted and rhizome elongation halved. Similar effects were elicited by indole-3-acetic acid, 5 × 10^-5, M in seawater and by sap expressed from C. racemosa or C. sertularioides and added to seawater at 2.5–10% by volume. The regulation of development in an algal coenocyte is discussed and analogies with regulation in multicellular plants are drawn.     

Strong regeneration ability from rhizome fragments in two invasive clonal plants (Solidagocanadensis and S. gigantea)

The two rhizomatous perennials Solidago canadensis and S. gigantea belong to the most widespread alien plants in Europe. Anecdotal observations suggest that they disperse also by rhizome fragments. For testing their resprouting ability, rhizome fragments of different sizes from both species were buried at four different soil depths (0, 5, 10 and 20 cm, respectively) in a common garden. Rhizome fragments of S. canadensis ranged 3–6 cm in length, those of S. gigantea 5–20 cm in length. Resprouting plants were harvested after 3 months and growth related traits measured to assess their vitality. Resprouting rate in S. gigantea was far higher than in S. canadensis (85 and 19%, respectively). In S. gigantea, fragments of all sizes resprouted from all soil depths whereas none rhizome of S. canadensis emerged from 20 cm burial depth. In S. gigantea, size related traits showed significant interactions between fragment size and burial depth, but not relative shoot growth rate. At all burial depths, vitality of plants emerging from small rhizomes was lower than plants emerging from large rhizomes. Effects of rhizome size became stronger with increasing burial depth. The results show that both species are able to resprout from buried rhizome fragments, and that succesful regeneration is more likely to occur in S. gigantea. Managing these species should avoid any activities promoting rhizome fragmentation and dispersal of fragments.     

Nonlinear processes in seagrass colonisation explained by simple clonal growth rules

The development of single clones of the seagrass Cymodocea nodosa was analysed using a growth model based on the formation of structures limited by diffusive aggregation. The model implemented the measured clonal growth rules (i.e. rhizome elongation and branching rates, branching angle, and spacer length between consecutive shoots) and shoot mortality rate for C. nodosa at Alfacs Bay (Spain). The simulated patches increased their size nonlinearly with time displaying two different domains of growth. Young patches showed a rapid increase with time of the length of rhizome network and the number of living shoots, which depended on rhizome branching rate, and increased the radial patch size (R_g) algebraically with the number of living shoots as R_g ∝ N_s^1/Df, being D_f the fractal dimension of the patch structure. Patches older than 4 years increased the production of rhizome network and the number of living shoots much more slowly, while their radial patch size behaved as R_g ∝ N_s^0.5 resulting from an internal patch compactation. Moreover, the linear growth rate of the simulated patches changed up to 30 fold during patch development, increasing with increasing patch size until patches reached an intermediate size. The modelled patch development was found to closely reproduce the observed patch structure for the species at the Alfacs Bay (Spain). Hence, the growth of C. nodosa patches initially proceeds with a growth mode controlled by the branching pattern (branching frequency and angle) of the species, producing sparse and elongated patches. Once patches exceed 4–5 years of age and contained >500 shoots, becoming dense and circular, they shifts to a growth model typical of compact structures. These results explain previously unaccounted evidence of the emergence of nonlinear patch growth from simple clonal growth rules, and highlight the importance of branching frequency and angles as critical determinants of the space occupation rate of seagrasses and probably other clonal plants.     

扎龙湿地不同生境芦苇种群根茎数量特征及动态

采用单位土体取样,计测长度和生物量的调查与统计方法,对扎龙湿地保护区4个生境芦苇种群根茎数量特征进行比较分析。结果表明,芦苇5月10日左右返青后进入营养生长期,根茎长度6—8月份缓慢增加,8—10月份显著增加,后期是前期的3.5—10.3倍,生长季中后期是种群新根茎补充和生长的主要时期,不仅实现了种群空间扩展,并为营养繁殖储备更多繁殖芽;根茎生物量和干物质贮量6—8月份逐渐减少,8—10月份又逐渐增加,均以生长季末期的10月份最大,并均显著地(P0.05)高于其他月份,种群根茎养分的消耗主要供给根茎芽的萌发和幼株生长,根茎养分的储藏又为翌年种群的更新及扩展提供物质保障,种群对地下根茎存在明显的养分"超补偿性"贮藏现象。种群根茎长度和生物量均以湿生生境最大,依次为旱生生境、水生生境,盐碱生境最小,根茎干物质贮量以旱生生境最大,依次为湿生生境、水生生境,盐碱生境最小。种群根茎长度与返青后实际生长时间之间均较好地符合直线函数关系,种群根茎生物量和干物质贮量与生长时间之间较好地符合二次曲线函数关系,R2在0.804—0.997之间,拟合方程均达到了显著或极显著(P0.01)水平。4个生境芦苇种群在根茎长度、生物量、干物质贮量等数量特征均表现出由遗传因素控制的比较稳定的季节动态规律,在生境间的差异及其差异序位又均基本稳定,均表现出明显的土壤因子环境效应,其中土壤含水量、有机质、速效氮为正向驱动,p H、速效磷为负向驱动,土壤含水量、p H对根茎数量特征的驱动作用更突出。     

Ediacaran macroalgal holdfasts and their evolution: a case study from China

A macroalgal holdfast (root-like structure) anchored or grown into sediments is a key trait of metaphytes and eukaryotic algae. Various patterns and taphonomic variants of congeneric holdfasts are preserved on the bedding planes of black shales of the Ediacaran Wenghui biota in South China. The macroalgal holdfast, which commonly consists of a rhizome, rhizoid and pith (perhaps mechanical tissue), can be morphologically classified into ten types within four groups of rhizomes: bare rhizome holdfasts (Grypania, Tongrenphyton and Sectoralga-type holdfasts), canopy rhizome holdfasts (Gemmaphyton, Gesinella, Discusphyton and Baculiphyca-type holdfasts), pithy rhizome holdfasts (Zhongbaodaophyton and pithy-cone-type holdfasts) and differentiated rhizome holdfasts (Wenghuiphyton-type holdfasts). Analysis of the Precambrian macroalgal record indicates that rhizomes played a more important role in the evolution of macroalgal holdfast than rhizoids. The following evolutionary stages of development of macroalgal holdfasts are proposed: (1) growth of the basal thallus into sediments (Grypania-type holdfast); (2) development of a primitive (indistinct) rhizome from the base of a stipe or thallus (Tongrenphyton-type and Sectoralga-type holdfasts); (3) growth of a distinct rhizome with rhizoid (Gemmaphyton-type, Gesinella-type and Discusphyton-type holdfasts); (4) development of a pith in the stipe (Baculiphyca-type holdfast); (5) pith extension into the rhizome (Zhongbaodaophyton-type and pithy-cone-type holdfasts); and (6) rhizome differentiation and development of a complex holdfast system (Wenghuiphyton-type holdfast).     

REGIONS OF CELL DIVISION AND ELONGATION DURING STEM GROWTH OF XANTHIUM

Regions of cell division and cell elongation were established during stem growth of Xanthium pensylvanicum (cocklebur). From percent mitosis it was determined that the region of cell division in a Xanthium stem of Plastochron Index (PI) 13.89 was 20 mm long, starting from the stem apex and proceeding in a basipetal direction. Measurements of cortical cell lengths demonstrated that mature cell length in the stem of the same plant was reached at about 55 mm distance from the stem apex. Between 20 and 55 mm distance from the apex, shoot growth occurred by cell elongation alone. The maximum rate of change in cell length (dC/dX) was at about 15 mm distance from the stem apex. The length of the apical growth region was a function of the age of the plants. The older the plant, the longer its apical region of elongation. Growth of Xanthium stems was due primarily to elongation of internodes; the nodal regions did not seem to elongate.     

STUDIES ON THE GERMINATION OF SEEDS OF THE ROOT PARASITES,ALECTRA VOGELII AND STRIGA GESNERIOIDES. I. ANATOMICAL CHANGES IN THE EMBRYOS

Anatomical changes in the radicle and shoot meristems of embryos of germinating seeds of the obligate root parasites, Alectra vogelii and Striga gesnerioides were studied. When germination of seeds was stimulated by cowpea (Vigna unguiculata) root exudate, growth occurred mainly in the radicular pole of embryos and minimally in the plumular pole, resulting in seedlings with elongated radicles. Maximum radicle elongations of about 3 mm in A. vogelii and 2 mm in S. gesnerioides were recorded during a period of 8 and 11 days, respectively. Analysis of the radicular tip during the course of seed germination revealed that the activity of the meristematic tissue progressively decreased until it completely disappeared. When germinated seeds were cultured on nutrient agar media, the radicle meristem of A. vogelii continued to grow producing a normal root with a root cap. On the other hand, the radicles of cultured S. gesnerioides seeds elongated only slightly before meristematic activity ceased. During continued growth of seedlings of both species on agar media, lateral roots whose tips had typical angiosperm root topography, were initiated from the radicle.     

Production of rhizoids by Caulerpa prolifera in culture

Studies on Caulerpa prolifera rhizoids were carried out to determine the possibility of mass culture, because the rhizoids produce a bio-adhesive. Rhizoids can be induced by cutting the base of a blade and floating it in a media or planting it in sand. Measurement of rhizoid production included determination of number, length, and the weight of attached sand grains. The growth experiments were for 1–2 weeks and fronds growth was compared to rhizoid production. Optimal conditions for rhizoid growth included low levels of nitrogen and phosphate (less than 5 and 2 μM, respectively), low irradiance (30 μmol photon m⁻² s⁻¹), moderate temperature (22–28°), continuous shaking, addition of microelements and auxin (1 ppm) and initially detached fronds followed by attachment. Under these optimal conditions maximal weekly growth reached 70–170 rhizoids per blade, 7–11 mm length and maximal attachment to sand grains. Blade growth of C. prolifera responded similarly to rhizoid production and reached a weekly growth rate of 30–130%.     

Effects of continuous illumination on the rhizome of Caulerpa prolifera

Caulerpa plants were grown under a 12-hr light and 12-hr dark(12L-12D) regime for 8 days followed by 8 days of continuouslight (24L-0D). Under both light regimes the elongation of therhizome was by means of "tip growth". However, the rate of rhizomeelongation in 24L-0D regime (10.9 mm/day) was higher than thatof 12L-12D regime (7.9 mm/day). Jn the region of 1.5 mm fromthe tip, the RERE (relative elemental rate of elongation) under24L-0D and 12L-12D regimes were respectively 4% and 3% elongationper hr. Under 12L-12D photoperiod the subapical part of therhizome exhibited distinctive oscillation: bending upward duringthe light period or at the time of rhizoid cluster initiation,and in the dark becoming relatively straight except at the timeof cluster initiation. No such distinctive oscillation was observedin continuous light. ¹ Report of work supported by research grant GB-7885 from theNational Science Foundation, and in part by Research Councilof Rutgers University. (Received June 8, 1971; )     

Negative Phototropism in the Rhizoid of Bryopsis plumosa

Negative phototropism in the rhizoid of Bryopsis plumosa wasanalyzed. The growth zone of the rhizoid was confined to theapical hemisphere, as is typical of tip growth. Upon unilateralillumination, the rhizoid bent away from the light source witha "bulging" manner. The photoreceptive site for phototropismwas also restricted to the apical hemisphere. The action spectrumfor this negative phototropism was determined from fluence-responsecurves that were obtained after fixing the duration of illuminationat 60 min and varying the fluence rate between 0.1 and 3.0 Wm^-2. The action spectrum had a large peak at 467 nm and smallerpeaks at 378 and 414 nm, resembling the action spectra of certainother blue-light responses. (Received November 29, 1995; Accepted May 29, 1995)