Storage carbohydrate production and overwintering strategy in a winter-green tussock grass on South Georgia (Sub Antarctic)

Summary Water-soluble and ethanol-soluble carbohydrates were extracted from leaves, shoots, roots, rhizomes and flowers ofParodiochloa flabellata (Lam.) Rasp. (=Poa flabellata (Lam.) Hook f.) throughout a growing season. Very high levels of soluble carbohydrates were found in the shoot, comprising up to 71% of the shoot dry weight. There was no significant storage in the rhizome. It is suggested that the production of storage fructans with a degree of polymerization greater than 14, rather than oligosaccharides or starch, may be related to the cold environment. Relationships between the onset of senescence and levels of fructans suggest that recovery of sugars from dying tissue is high. The sugar reserves together with preformation of the flowers should ensure production of seed however poor the summer or long the winter might be. The high levels of fructans may partly inhibit growth but may also offer some degree of freezing resistance.     

Growth forms of Leymus chinesis (Poaceae) at the different developmental stages of the natural population

The manner in which the density of Leymus chinensis increases from a single plant to a dominant population can be understood by tracing the development of a population from early to late stages. Parent shoot density, above‐ground dry weight, spike density, heading rate and spike dry weight, density of spreading shoots (buds/daughter shoots in apical/axillary rhizomes) and clumping shoots (buds/daughter shoots in axillary parent shoots), and young rhizome length and weight were investigated in the same quadrats for a low density/early stage (LE) population and a high density/late stage (HL) population. Clonal growth (buds/daughter shoots formation) and sexual reproduction (spikes formation) increased while rhizome storage (young rhizome weight) decreased during the transition from LE to HL. In a LE population an outward occupation strategy was employed, with a high proportion of spreading shoots. As the population density gradually increased until HL, an inward consolidation strategy increasing shoot amount in previously occupied areas, was adopted. This was characterized by a high proportion of clumping shoots. Interestingly, the trade‐off between spreading and clumping shoots can be adjusted by the duration of young rhizome elongation during a growth season. In other words, compared with a HL population, a LE population shortened the duration of young rhizome elongation during the growth season, which resulted in more time for the production of axillary spreading shoots along the rhizomes, and high amounts and proportions of total spreading shoots. The special growth patterns, that is, trade‐offs among growth forms, allow L. chinensis to establish dominant populations throughout the eastern Eurasian Steppe.     

Life history trade-offs in Amphibromus scabrivalvis (Poaceae): allocation to clonal growth,storage, and cleistogamous reproduction

Life history trade-offs among clonal growth, storage, and sexual reproduction were investigated in the perennial grass Amphibromus scabrivalvis in relation to soil nutrients. This species exhibits clonal growth by producing rhizomes and stores reserves in the form of basal corms; seeds are matured in cleistogamous spikelets on panicles enclosed within the leaf sheaths along each culm. Ten seed-derived genotypes (clones) were separated into 72 ramets and planted in the greenhouse. Control ramets received only water while the remainder received fertilizer applied every 2 wk. Twenty-four ramets were harvested per clone at 11, 20, and 26 wk. The dry mass of corms, rhizomes, roots, shoots, and seeds were recorded. Biomass partitioning to rhizomes provided a measure of carbon allocation to clonal growth, partitioning to corms provided a measure of allocation to storage, and partitioning to seeds provided a measure of allocation to sexual reproduction. Allocation to most organs was significantly influenced by clone identity; fertilizer significantly increased allocation to corms and seeds at 20 wk, but never affected rhizome allocation at any age. Corm allocation increased from 2% at 11 wk to 27% at 26 wk; rhizome allocation decreased from 10% at 11 wk to 3% at 26 wk. Significant negative relationships were detected for rhizome vs. seed and corm vs. rhizome allocation in fertilized clones at 20 wk. This suggests an age-dependent physiological life history trade-off between clonality and sexual reproduction and between clonality and storage. In contrast, a significant positive relationship was consistently noted for corm vs. seed allocation in fertilized and unfertilized clones at 20 and 26 wk. The absence of a trade-off between storage and sexual reproduction may indicate that these two processes are not necessarily mutually exclusive components of life history.     

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.     

Seasonal patterns of partitioning and remobilization of 14C in the invasive rhizomatous perennial Japanese knotweed (Fallopia japonica (Houtt.) Ronse Decraene)

Resource partitioning between shoot growth, storage and reproduction is poorly understood in many clonal plant species. This study documents seasonal patterns of growth, ¹⁴C-labelled photoassimilate distribution and remobilization in the invasive rhizomatous species Fallopia japonica (Japanese knotweed). Biomass accumulation above- and below-ground in F. japonica was rapid. By September, rhizome biomass had increased 18-fold from the initial harvest in May (representing 48% of total plant biomass) and this was maintained over winter. Patterns of ¹⁴C allocation from F. japonica shoots labelled at different times of year show that as the season progressed, the rhizomes became an increasingly important sink for current assimilate (the percentage of ¹⁴C recovered from rhizomes was 35% in August and 67% in September) and the corresponding retention of assimilate by established shoots declined. The percentage of ¹⁴C exported to roots was greatest in August. Relatively little photoassimilate was exported to other shoots on the plant, or to flowers. Recycling of photoassimilate was fairly tight in this species and ¹⁴C fixed by shoots in early May 1999 or September 1999 was remobilized to the rhizome prior to shoot senescence and death. Some of this ¹⁴C was then remobilized to new shoots early the following spring. These characteristics may contribute to the success of F. japonica in colonizing a variety of contrasting habitats, often with serious management implications.     

In vitro propagation and field establishment of Eulophia cullenii (Wight) Bl., a critically endangered orchid of Western Ghats, India through culture of seeds and axenic seedling-derived rhizomes

An efficient protocol has been devised for the propagation and field establishment of Eulophia cullenii (Wight) Bl., a terrestrial orchid having ornamental potentialities, and is critically endangered in Western Ghats, India. Seeds extracted from 60–90-d-old capsules germinated in ½ MS, ¼ MS, Knudson C, or Mitra liquid medium developed into 1.4–2.5-mm-diameter protocorms in 60 d. Supplementation of organic additives like coconut water, peptone, yeast extract, and casein acid hydrolysate (CH) significantly enhanced protocorm growth. Upon subculture onto agar-gelled Mitra medium fortified with 0.05% CH, 56% of protocorms regenerated into shoots through the formation of linear mini-rhizomes. The regenerated shoots grew vigorously in ½ MS, producing new rhizomes. Mature rhizomes from axenic seedlings produced maximum (13?±?1.4) shoots/whole rhizome in ½ MS fortified with 44.4 μM 6-benzylaminopurine (BAP), in 120–150 d. Horizontal and longitudinal halves of the rhizome also gave multiple shoots (6–8.5) in the presence of 44.4 μM BAP. Shoots or shoot clumps sub-cultured onto ½ MS basal medium produced roots followed by rhizomes in 60–150 d. Seedlings with mature rhizomes showed 70% establishment in the nursery and added a new rhizome at the end of one growth cycle. An average of 70.6% of the rhizomes originating from seedlings during the second growth cycle sprouted to produce new shoots, when planted in the native localities. Asymbiotic germination and cloning through rhizomes thus can provide a large number of vigorous plants of E. cullenii for ornamental exploitation as well as eco-restoration, if rhizome as storage organ is ensured in the propagule.     

Localization of lily symptomless virus and cucumber mosaic virus in anther- and filament-derived calluses and effect of callus culture duration on virus-free bulblet production in Lilium × ‘Enchantment’

Bamboos represent one of the world’s great natural and renewable resources. The study reports precocious rhizome formation in multiple shoots of elite, rare, woody bamboo Bambusa bambos var. gigantea. Multiple shoots were initiated from embryonic axes of caryopses inoculated on MS-basal medium supplemented with 5.0 μM BAP and 2% sucrose. Transfer of shoots to MS basal medium supplemented with BAP (2.5 μM, 5.0 μM), GA₃ (0.1 μM) and NAA (50.0 μM) and 5% sucrose led to 58% to 100% rhizome induction within four weeks of culture. Subsequently, these rhizomes developed roots on auxin media and formed culm shoots showing regeneration of plantlets after eight weeks. Incorporation of TIBA inhibited rhizome formation. The plantlets with rhizomes were transferred to soil. Precocious rhizome formation will lead to early establishment besides providing propagules on demand and mass multiplication of bamboos through rhizome banks.     

Seasonal changes in growth and metabolic activity of underground shoots of yarrow

Growth and metabolic activity of underground shoots of a long-rhizome perennial herbaceous species yarrow (Achillea millefolium L.) were studied. The active growth of rhizomes and the formation of new meristematic zones were observed during the second half of the growing season after termination of aboveground shoot growth. In this period, underground shoots had a rather high rate of respiration (1.3 mg CO₂/(g dry wt h)), a considerable content of nonstructural carbohydrates (15% of dry weight), and the elevated activities of IAA, cytokinins, and ABA. In autumn, the rate of respiration of underground shoots decreased to 1.0 mg CO₂/(g dry wt h), soluble sugars accumulated, the ratio between unsaturated and saturated fatty acids rose as well as the ratios GA/ABA and cytokinins/ABA. Temperature optimum for the rhizome growth lay in the range of low and moderate above-zero temperatures (5–20°C), and the freezing point of water in the apices of under-ground shoots was about ?10°C. It is concluded that rhizome quiescence predominantly depends on low temperatures and is not associated with the accumulation of growth inhibitors. In the course of plant preparation to winter, morphogenetic transformations in underground shoots depend on changes in the hormonal balance directed in favor of growth hormones and relatively high respiratory activity in the apical zones of the rhizomes.     

TRANSLOCATION IN POLYTRICHUM COMMUNE (BRYOPHYTA). II. CLONAL INTEGRATION

Polytrichum commune is a perennial moss that grows by rhizomatous production of interconnected shoots. It possesses a well-developed internal system of food-conducting tissue. Liquid scintillation determinations following pulse-¹⁴CO₂ treatment of a single shoot in rhizomatously connected groups indicate that tagged photoassimilates are transported into:1) newly developing stems during the early growing season; 2) shared rhizomes (with an increase observed in rhizome allocation toward the end of the year, as well as a year-end percentage increase in starch reserves); and 3) mature neighboring shoots. Over the course of the growing season, photoassimilate export from pulse-labeled shoots ranges between 12.9% and 21.4% of total tagged. The amount of export is significantly influenced by defoliation, shading, stem tip removal, and 50 ppm indole-3-acetic acid treatments performed on rhizomatously connected neighbor shoots. Physiological integration within P. commune can be inferred from these results. Control is exerted by endogenous hormones or by changes in the activity of sources and sinks.     

扎龙湿地芦苇种群不同龄级根茎构件的季节动态

克隆植物根茎具有营养繁殖和扩展种群的功能,也是芽和分株生理整合的通道。根茎构件具有出生、死亡及年龄等种群统计特征,不同龄级根茎的季节动态可以反映根茎的存活和衰老过程。采用单位土体挖掘取样,对扎龙湿地4个生境芦苇种群根茎构件进行野外调查,比较不同龄级根茎长度、生物量和干物质贮量的季节动态。结果表明:7—10月份,1a根茎长度、生物量和干物质贮量均呈指数函数增加,在生长季中后期有一个持续时间较长的生长和物质积累时期。6—10月份,2a、3a根茎长度呈线性函数增加,4—6a根茎长度呈线性函数减少;2—4a根茎生物量和2—5a根茎干物质贮量呈二次函数先减少后增加,5a、6a根茎生物量和6a根茎干物质贮量呈幂函数减少。整个生长期内,根茎长度和根茎生物量均以3a最大,根茎长度以最高的6a最小,根茎生物量以最低的1a最小;根茎干物质储量以5a最大,以最低的1a最小。4个生境芦苇种群根茎长度、生物量和干物质贮量在龄级间的差异及差异序位稳定,在新根茎的产生、老根茎的存活以及根茎寿命与养分消耗和储藏上均具有稳定的生物学特性,不同龄级根茎在种群中的地位和作用以及对种群的贡献不同。     

Rhizome development ofPhragmites australis in a reed community

Variations in the height, shoot density, biomasses of above- and below-ground parts and rhizome distributions ofPhragmites australis were investigated along a line-transect in a reed community at Yufutsu Mire, Hokkaido. Relationships of performance of the reed plants to soil conditions and species compositions were also examined. Three types of rhizome development were recognized in reed plants; (1) the central part of the reed community, characterized by well developed rhizomes and dense aerial shoots, (2) the intermediate part, characterized by development of rhizomes along both the peat and surface layers and very dense aerial shoots, and (3) the marginal part, characterized by development of rhizomes only along the peat layer and sparse aerial shoots. Observation showed that rhizomes in the surface layer actively produced aerial shoots, whereas rhizomes in the peat layer contributed to the spreading of their distribution range. With the growth of rhizomes, organic debris originating from dead rhizomes and roots accumulated in the mineral soil to promote organic soil formation. In dense parts of the reed stand, species composition was poor because of the shading and litter accumulation by reed plants. The effects of microtopography and water level on the establishment of reed seedlings were also considered.     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Seasonal allocation of carbohydrates between above‐ and below‐ground organs of Typha domingensis

Seasonal carbohydrates allocation by Typha domingensis was evaluated to identify the potential physiological weaknesses in the growth cycle of this plant in Lake Burullus, Egypt. Monthly plant samples (February–October 2014) were separated into shoots, roots and rhizomes to evaluate the seasonal changes in water‐soluble carbohydrates (WSC), starch and total non‐structural carbohydrates (TNC) for each plant organ. The present study indicated that rhizomes are strong carbohydrates sink during the life cycle of T. domingensis. Starch represented the greatest part of the TNC pool, surpassing the concentration of WSC 1.8–4.3 times. The WSC, starch and TNC concentrations of T. domingensis below‐ground organs (rhizomes and roots) were high at the beginning of the vegetative period (February); they reached their minima in March to support the shoots growth, then were followed by a gradual increase due to the translocation from shoots. The time when T. domingensis is expected to be most susceptible to a management technique is at the point in the seasonal cycle when the stored carbohydrates are at the lowest (in March).     

Effects of lead contamination on the clonal propagative ability of Phragmites australis (common reed) grown in wet and dry environments

Clonal propagation is important for the survival and maintenance of the common reed Phragmites australis. Pot culture experiments were conducted to investigate the effects of lead (Pb) concentration (0, 500, 1500, 3000, 4500 mg·kg^?1) and water stress on the clonal reproductive ability of this species. The Pb concentration found in plant organs, in decreasing order, was roots >shoots >rhizomes. There was a negative relationship between the growth of clonal propagative modules (excluding axillary shoot buds) and Pb concentrations, which caused a decrease in biomass, rhizome growth and number of axillary and apical rhizome buds. Daughter axillary shoots exhibited a tolerance strategy, with no significant change in their number; the axillary and apical rhizome buds, daughter apical rhizome shoots and rhizomes exhibited compensatory growth during the late stage of Pb (excluding 4500 mg·kg^?1) treatment in a wet environment. Pb applications above 500 mg·kg^?1 reduced these parameters significantly in the drought treatment, except for the number of axillary shoot buds, which did not change. Our results indicate that clonal propagative resistance to Pb contamination can occur via tolerance strategies, compensatory growth and a Pb allocation strategy, enabling these reeds to maintain population stability in wet environments. However, clonal modular growth and reproductive ability were inhibited significantly by the interaction between drought and Pb, which would cause a decline in P. australis populations in a dry environment. Lead concentrations of 4500 and 500 mg·kg^?1 in soils might meet or exceed the Pb tolerance threshold of clonally propagated reeds in wet and dry environments, respectively.     

Storage and mobilization of nonstructural carbohydrates and biomass development of beech seedlings (Fagus sylvatica L .) under different light regimes

 Beech seedlings were grown under 8%, 13%, 23% and 100% relative light intensity for 2 years after germination. Starch, sucrose and monosaccharides from the bark and wood parenchyma of shoots and roots were analyzed during the course of the second year. The annual allocation pattern of starch revealed five successive phases: starch disintegration in November (1) was paralleled by high monosaccharide concentrations in the shoot cortex (≤ 33.4 mg/g DW). Seedlings of all light variants reached maximum sucrose concentrations (≤ 82.8 mg/g DW) during starch disintegration in January (2) that coincided with decreased monosaccharide contents. Up to mid-April, resynthesis of starch (3) occurred in most shaded and unshaded seedlings. In May, starch was converted into monosaccharides in all storage tissues (4). Seedlings grown under 13% light intensity showed de novo synthesis of starch (5) 4 weeks after bud burst. These seedlings reached 98% maximum starch storage capacity of the shoot and 89% of the root in July. In mid-October, the maximum starch concentration of the roots increased with light intensity, and this corresponded with an increase of lateral root growth. The variation of shoot and root dry weight was closely related to the content of nonstructural carbohydrates during the second year. The shift of shoot growth to the first half of the growing season and the suppression of lateral root growth during the second half is assumed to be a strategy of young beech to survive under light limiting conditions. Received: 22 May 1997 / Accepted: 10 October 1997     

Integrated growth patterns of turtle grass,Thalassia testudinum Banks ex König

Indications of physiological integration in the clonal plant Thalassia testudinum Banks ex König from the Puerto Morelos reef lagoon, Mexican Caribbean, were deduced from synchronisation in the formation of ‘inactive shoots’, lateral rhizomes and inflorescences. ‘Inactive shoots’ (i.e. bare, pointed shoots with live roots attached) on a rhizome section often had similar numbers of leaf scars. Lateral rhizome sections were generally found in similar positions when a rhizome had more than one shoot bearing a lateral rhizome. Additionally, the position of lateral rhizomes and the number of leaf scars on inactive shoots were often similar when encountered on the same rhizome section. Synchronisation of flowering events was suggested by the similar position of inflorescence scars on different shoots on the same rhizome section. It is suggested that inactive shoots can play a role in density regulation of the clonal population, and that they possibly constitute a ‘dormant meristem bank’ analogous to ‘seed banks’, or ‘dormant bud banks’.     

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。     

Demography of Carex brevicuspis (Cyperaceae) rhizome populations: a wetland sedge that produces both elongated and shortened rhizomes

The wetland sedge Carex brevicuspis reproduces vegetatively by producing short rhizomes to form clumping ramets phalanx) and long rhizomes to form spreading ramets (guerrilla), resulting in a combined growth form. As an initial step towards understanding the adaptation of Carex growth strategies to seasonal fluctuations in wetland habitats, we investigated the density and composition of C. brevicuspis rhizome populations immediately after flooding (November), in winter (January), in spring (March), and before flooding (May) in the Dongting Lake wetlands, China. The total rhizome density peaked in winter and was lowest before flooding. A large rhizome population in winter may enable C. brevicuspis to survive the seasonal cold weather and recruit a shoot population in the spring. A small rhizome population before flooding may optimize reproductive allocations and be a strategy for enduring the long flooding season. Regardless of date, short rhizomes comprised the majority of the rhizome population (73.0% in March to 98.2% in May). This indicates that C. brevicuspis primarily uses a phalanx growth strategy to utilize locally abundant resources in wetlands. The percentage of long rhizomes in the rhizome population varies significantly between seasons (1.8% in May to 27.0% in March), indicating that growth form also changes with seasonal fluctuation of wetland habitats. The results show that C. brevicuspis may adapt to seasonal changes in wetland habitats through changes in demography of rhizome populations.     

Vegetative propagation and sexual reproduction in the woodland understorey pseudo-annual Circaea lutetiana L.

On the forest floor of deciduous woodlands, relative light intensity gradually declines during the early growing season. The woodland understorey pseudo-annual Circaea lutetiana L. completes its life-cycle at the end of summer. These pseudo-annuals are clonal plants which survive the winter only as seeds and as hibernacles produced by the rhizome apices. In this paper, we asked several questions related to the life-history of C. lutetiana. It was found that shoot formation in early spring did not exhaust the old hibernacle. A trade-off between rhizome + hibernacle number and weight might be expected when plants grow under resource limitation. It was hypothesised that both number and weight of rhizomes and hibernacles will be affected by light availability. Since the effect of resource supply on the size number trade-off will depend on the developmental pattern of the rhizome system, rhizome development was studied as well. Soon after the shoots emerged, 1st order rhizomes were formed in May on the nodes of the old hibernacle. First-order rhizomes branched in June and 2nd order rhizomes (side-branches) were continuously produced throughout the growing season. The phenology and developmental rate of plants growing in different light treatments were plastic. On average 30% of rhizome biomass was formed during the vegetative phase, and rhizome and fruit production were only partly separated in time. The ratio of total rhizome biomass to total fruit biomass was not affected by light. Also flower bud removal did not lead to an increase in rhizome production, which suggests that division of biomass to both reproductive modes is rather rigid. The number of 1st order rhizomes was not affected by the light treatments. Under light limitation, both rhizome number and weight of single rhizomes were reduced. In contrast, fruit number, but not weight of single fruits, was limited.     

RHIZOME AND ROOT GROWTH RATES AND CYCLES IN PROTEIN AND CARBOHYDRATE CONCENTRATIONS IN GEORGIA SPARTINA ALTERNIFLORA LOISEL. PLANTS

Root and rhizome growth and composition were measured in a Georgia salt marsh. Root growth was less seasonal than rhizome growth; none of the latter occurred during the first quarter of the year. The depth of underground growth shifted downward during the fall. Sugar and starch concentrations were relatively low in roots throughout the year. They remained relatively high in the rhizomes, but varied in distribution depending on the season.