A modelling approach to the analysis of pressure-flow in Phragmites stands

Field studies have shown that resistance to convective gas-flow within the culm-rhizome gas-space system of Phragmites tends to be higher in ‘die-back’ as opposed to healthy stands. However, the collection of reliable data and its interpretation is complicated by many factors. To help interpret field measurements and further our understanding of convective flows in general, we have developed mathematical models based on the humidity-induced convective-flow generating potential of culms. Among other things, these make it possible to study the effects on pressure flows of increasing flow resistances in rhizomes and between culms, of different pressure-generating potentials of interlinked culms and of different numbers of efflux culms. A multi-culm and rhizome model is described together with some simple examples of the way it can prove helpful in interpreting some of the field observations from healthy and die-back sites.Increased venting resistance reduced flows curvilinearly; increasing the counter pressure to mimic those induced by other interlinked living culms reduced flows linearly. In some of the multi-culm examples shown counter-pressure exerted an effect approximately the same as that of rhizome and venting resistance, but as culm numbers declined it assumed even greater importance. The value of conductivity derived from applying pressures to the stubble of excised culms and measuring the flows induced, proved to be a composite measure of the effective conductivity of the whole rhizome-culm train rather than that of the rhizome plus major vent. The expression 1−(P_d/P_s) — the delivery coefficient — where P_d is the dynamic pressure in the base of the intact culm, and P_s the static pressure developed by the culm with the outflow blocked, was identified as a useful and easily obtained measure of the ability of an individual intact culm to contribute to the convective gas-flow in a stand, and one which should be relatively unaffected by weather conditions. For identical interlinked culms it was shown that the flows from all culms fitted exactly along the same line of declining flow versus dynamic pressure irrespective of the numbers of culms or number and position of outflow vents.     

Density regulation during the regeneration of two monocarpic bamboos: self-thinning or intraclonal regulation?

Abstract. The population dynamics of two monocarpic bamboos, Sasa kurilensis and S. tsuboiana, were studied for more than 10 years after establishment following mass flowering. Both species show vigorous rhizomatous vegetative reproduction after growing up to maturity, but horizontal expansion in the seedling stage was much more vigorous in S. tsuboiana than in S. kurilensis. The pattern of changes in culm density in the two species was strikingly similar: culm densities of both species increased until they reached full-density states, after which they decreased in accordance with seedling growth. However, the mode of regulation in culm density was different. S. kurilensis seedlings were composed of only a few culms and scarcely extended their rhizomes during the observation period. Such poor lateral expansion resulted in asymmetric competition as observed in many non-clonal plants, and consequently their culm density decreased as a result of the mortality of genets due to self-thinning. In S. tsuboiana seedlings, the number of culms per genet increased considerably by frequent tillering and sprouting from rhizomes. However, after reaching full density state, the Bud Utility Ratio (BUR), (the proportion of the rhizome nodes with culms to the total number of rhizome nodes), decreased drastically. In this manner, S. tsuboiana regulated culm density intraclonally as is observed in the stable states of many clonal plants. Hence it is important for the understanding of the regeneration process in clonal species to clarify when and how their seedlings extend rhizomes during their growth.     

Temporal and spatial profiling of internode elongation-associated protein expression in rapidly growing culms of bamboo

In natural conditions, culms of developing Moso bamboo, Phyllostachys heterocycla var. pubescens, reach their final height of more than ten meters within a short period of two to four months. To study this phenomenon, bamboo culm material collected from different developmental stages and internodes was analyzed. Histological observations indicated that the development of culm was dominated by cell division in the initial stages and by cell elongation in the middle and late stages. Development, maturation, and aging in different regions of the culm were studied systematically from the basal to the top internode. The four major endogenous hormones, indole acetic acid, gibberellic acid, zeatin riboside, and abscisic acid appeared to strongly influence the cell elongation phase. A total of 258 spots were differentially expressed in culm development. Of these, 213 spots were identified by MALDI-TOF/TOF MS and were involved in many physiological and metabolic processes including carbohydrate metabolism, cell division, cell expansion, protein synthesis, amino acid metabolism and redox homeostasis. These proteins with different expression patterns constructed an ingenious network to regulate the culm development. Developmental stage-specific and internode-specific protein expression patterns were identified. Protein abundance was regulated temporally and to some extent spatially, and the sequential development from base to apex of bamboo culm was implemented by temporal and spatial expression of enzymes. Results indicate that during development energy was mainly derived from sucrose degradation, as photosynthetic capacity was poor. The regulation of anaerobic and aerobic modes of respiration appeared to play an important role in energy generation. This is the first report on proteomic profiling in bamboo and helps in understanding the regulatory processes in developing culms.     

Pressure gradients along whole culms and leaf sheaths, and other aspects of humidity-induced gas transport in Phragmites australis

Emergent aquatic macrophytes growing in waterlogged anaerobic sediments overlain by deep water require particularly efficient ventilating systems. In Phragmites australis (Cav.) Trin. ex Steud, pressurized gas flows, generated by humidity-induced diffusion of air into leaf sheaths, enhance oxygen transport to below-ground parts and aid in the removal of respiratory CO2 and sediment-generated CO2 and methane. Although modelling and flow measurements have pointed to the probable involvement of all leaf sheaths in the flow process and the development of pressure gradients along the whole lengths of living culm and leaf sheaths, direct measurements of pressure gradients have never been reported. The aim of this study was to search for pressure gradient development in Phragmites culms and leaf sheaths and to determine their magnitudes and distribution. In addition, dynamic (with gas flow) and static pressures (no flow condition) and their relationship to flows, leaf sheath areas, and living-to-dead culm ratios were further investigated. Dynamic pressures (DeltaPd) recorded in the pith cavities of intact (non-excised) leafy culms, pneumatically isolated from the below-ground parts and venting through an artificial bore-hole near the base, revealed a curvilinear gradient of pressure 'asymptoting' towards the tips of the culms. Similarly, DeltaPd in upper and lower parts of leaf sheaths increased with distance from the base of the culm, with values in the upper parts always being greater. Curvilinear gradients of pressure were also found along pneumatically isolated individual leaf sheaths, but radial channels linking the leaf sheath aerenchyma with the pith cavity of the culm appeared to offer little resistance to flow. In keeping with predictions, static pressure differentials (DeltaPs) achieved in intact and excised culms and single leaf sheaths on intact culms proved to be relatively independent of leaf sheath area, whereas the potential for developing convective flows (pressure-driven flows) increased with increasing leaf sheath area. As measured by the ventilating coefficient [1-(DeltaPd/(DeltaPs)] the old dead (efflux) to living (influx) culm ratio of 1:12 compared with 1:25 raised ventilating efficiency from 31% to 71%, giving flows per tall culm into the rhizome system of c. 2.8 cm3 and 6.5 cm3 min-1, respectively. It was concluded that dynamic pressure gradients probably extend along the whole length of the leafy culms and leaf sheaths of Phragmites and that all leaf sheaths and all exposed points along the leaf sheaths can contribute convective gas-flow to the rhizome system.     

水竹（Phyllostachys heteroclada）人工林生物量结构的研究

 根据39块标准地上测定18808株立竹,47个竹鞭样方,38株不同年龄(1—6年)和不同径级样竹,用相对生长量法(allometric method)研究不同水竹人工林地上和地下部分生物量结构;用数理统计方法拟合出竹秆、竹叶、竹枝和竹鞭生物量与胸径的回归方程。地上部分生物总量随径级而增加,竹秆重量按径级分布分别为,1.5径级与林分总重量比为16%;2.0径级为30.5%;2.5径级为26.1%,3.0径级为16.0%。地上部分生物总量随龄级增加而降低,分别为：Ⅰ龄级为31.7%,Ⅱ龄级为22.0%,Ⅲ龄级为17.5%,Ⅳ龄级以上为28.0%。竹鞭中壮鞭占50—60%,竹鞭容积2.0—2.5%,地上部分总量与地下总量比为1.0—1.5,叶面积指数为4—6,每亩产量可达4000—7000斤时,称为高产林分。     

Biology of the bamboo Chusquea culeou (Poaceae: Bambusoideae) in southern Argentina

Over a period of 7 years the biology and phenotypic variability of Chusquea culeou were studied at 5 locations in cool temperate forests of southern Argentina. Excavated rhizomes had an average of 1.1 successful rhizome buds, and an average of 2.1 years elapsed between successive generations of rhizomes. Rhizome buds usually develop within the first four years after a rhizome forms. Height, volume and weight of a culm can be calculated from its diameter 1 m above the ground. Culm size, length of foliage leaf blades, and pattern of secondary branching differed among study sites. Dead culms were numerous and commonly remained erect for more than 7 years after dying. New culm shoots appear in spring and reach full size within a few months. Shoots can grow more than 9 cm/day. Less than half of the shoots survived a year; most were killed by moth larvae. Multiple primary branch buds emerge through the culm leaf sheaths in the second spring. The mean number of branch buds at mid-culm nodes varied between 34.8 and 81.5, and the mean number of primary branches was between 22.8 and 40.8. Number and length of branches, and number and length of foliage leaf blades at each node is related to the position of the node on a culm. Most branches grow about 3 cm and produce 1 to 3 foliage leaves annually. Foliage leaf blades generally live 2 years or more; few survive 6 years. Relative lengths of foliage leaf blades and their spacing along a branch permit recognition of annual cohorts.Both gregarious and sporadic flowering have been reported, and every year a few isolated plants flower and die. Length of the life cycle is unknown. Seedlings require up to 15 years to produce culms of mature size. Foliage branches may live more than 23 years, and culms may survive 33 years. Extensive loss of new shoots to predation suggests that gregarious flowering may be driven by a need to escape parasitism. C. culeou clumps expand slowly. Average annual rate of increase of the number of live culms in a clump was 4.6%. Methods of seed dispersal are undocumented. A dense stand of Chusquea culeou had an estimated phytomass of 179 tons/hectare (dry weight), 28% of which was underground. Net annual production was about 16 t/ha dry weight.     

毛竹种群向针阔林扩张的根系形态可塑性

为了弄清毛竹(Phyllostachys edulis)向针阔林扩张过程中根系的形态可塑性反应,在浙江天目山自然保护区毛竹向针阔林扩张的典型过渡地带,连续区域上设置毛竹纯林、针阔-毛竹混交林(以下简称过渡林)、针阔林3种样地。用根钻法采集样地毛竹根系、针阔树根系并比对其生物量密度、细根比根长、相邻同级侧根节点距等形态特征参数变化。结果表明:随着毛竹的扩张程度增加,林内根系生物量密度增加;且与针阔树竞争过程中毛竹将更多的根系放置于表层;同时在水平方向上随离样株距离的增加未出现明显变化,而针阔树根系则随离样木距离的增加而逐渐减少;毛竹根系比根长明显增加,平均增幅15%;一、二级侧根节点距则均有所下降,毛竹侧根数量增多。这些结果表明毛竹种群可通过根系生物量密度、细根比根长、相邻同级侧根节点距等形态可塑性方式实现向周边森林扩张。     

Bolboschoenus (Ascherson) Palla (Cyperaceae) from Pliocene of China

Tubers, culms, rhizomes, and leaves of Bolboschoenus cf. yagara (Ohwi) YC Yang et M Zhan were reported from the sediments of Zhangcun Formation, Late Pliocene in Shanxi Province, China. The fossil species possesses trigonous culms. And tubers are cordiform, spindle shaped, or round, and are covered by roots in internodes and often connected by rhizomes. This is the first convincing fossil record of Cyperaceae in China.     

Bolboschoenus （Ascherson） Palla （Cyperaceae） from Pliocene of China

Tubers, culms, rhizomes, and leaves of Bolboschoenus cf. yagara (Ohwi) YC Yang et M Zhan were reported from the sediments of Zhangcun Formation, Late Pliocene in Shanxi Province, China. The fossil species possesses trigonous culms. And tubers are cordiform, spindle shaped, or round, and are covered by roots in internodes and often connected by rhizomes. This is the first convincing fossil record of Cyperaceae in China.     

Changes in abscisic acid and indole-3-acetic acid in axillary buds of Elytrigia repens released from apical dominance

Growth of axillary buds on the rhizomes of Elytrigia repens (L) Nevski is strongly dominated by the rhizome apex, by mechanisms which may involve endogenous hormones. We determined the distribution of indole-3-acetic acid (IAA) and abscisic acid (ABA) in rhizomes and measured (by gas-chromatography-mass spectrometry) their content in axillary buds after rhizomes were decapitated. The same measurements were also made in buds induced to sprout by removing their subtending scale leaves. The ABA content tended to be higher in the apical bud and in the axillary buds than in the adjacent internodes, and tended to decline basipetally in the internodes and scale leaves. IAA was similary distributed, except that there was less difference between the buds and other rhizome parts. After rhizomes were decapitated, the ABA content of the first axillary bud declined to 20% of that of control values within 24 h, while the IAA content showed no marked tendency to change. The ABA content also declined within 12 h in the first axillary bud after rhizomes were denuded, while the content of IAA tended to increase after 6 h. These changes occurred before the length of the first axillary bud increased 24–48 h after rhizomes were decapitated or denuded. We conclude that the release of axillary buds from apical dominance in E. repens does not require IAA content to be reduced, but is associated with reduced ABA content.     

神农箭竹开花前后植物体内N、P、K元素的动态

对神农箭竹(Fargesia murielae)N、P、K的含量进行分析,结果表明:N、P、K在竹子植物体内呈非均匀分布。N和P在各器官中的分布规律为:叶>鞭、根>竿;K在未开花竹和正开花竹中分布规律为:鞭>根>叶>竿,在已开花竹中分布规律为:叶>鞭>竿>根。随着开花过程的进行,N的含量在叶、竿、鞭和根等器官中逐渐减少;P的含量在竿、鞭和根等器官中逐渐降低,在叶片中先升高后降低;K的含量在鞭和根中呈下降趋势,在竿和叶中先降低后升高。     

Regeneration capacity and carbohydrate reserves in a clonal plant Rumex alpinus: effect of burial

The morphological and anatomical responses to different depths of burial were examined in Rumex alpinus (Polygonaceae), a perennial plant with monopodial, horizontally growing rhizome. Its segments, which consist of 12–20 internodes, 1 to 2 mm in length each, are products of single growing seasons. The rhizomes regenerated from 5, 10 and 20 cm, but failed to emerge from 30 cm. Number of internodes produced during a growing season was not affected by burial but the length of internodes increased up to about 30-fold. The rhizomes growing up to the surface were subsidized by older rhizome segments. In the case of deeply buried rhizomes the carbohydrate reserves of the last-year-segment were nearly completely depleted. Evolutionary significance of the regeneration capacity is discussed.     

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.     

Early Exposure to Ethylene Modifies Shoot Development and Increases Sucrose Accumulation Rate in Sugarcane

Sugarcane varieties (Saccharum spp. hybrids) that accumulate high levels of sucrose at the start of the harvest season are of considerable commercial interest. Our understanding of the factors that contribute to early sucrose accumulation in these varieties is limited. In this study we used the plant hormone ethylene to investigate the relationship between growth and early sucrose accumulation in sugarcane. The sugarcane variety KQ228 was exposed to a low concentration of the ethylene-forming compound 2-chloroethylphosphonic acid (CEPA) for a prolonged duration commencing from shoot emergence. The changes in sucrose accumulation and plant growth were investigated. Results from two glasshouse experiments revealed that the CEPA-treated plants accumulated a significantly higher amount of sucrose in their primary culm 2 and 3½ months post-germination. The treated plants had taller primary culms with many smaller internodes, smaller leaves, and a higher photosynthetic rate. Despite producing smaller internodes, treated culms were comparable in fresh weight and volume to the controls due to the compensating effect of faster internode formation. We identified three factors that may have contributed to the early accumulation of more sucrose in the treated culm: (1) the specific leaf area of young leaves was greater indicating efficient diversion of photoassimilate to sink tissue, (2) internode formation was initiated earlier, and (3) internodes continued to form at a faster rate. Consequently, a greater proportion of the internodes in the treated sugarcane matured earlier and began filling with sucrose sooner. The higher reducing sugar level in the apical region of the culm probably contributed to faster internode development. This coincided with elevated vacuolar and cell wall acid invertase gene expression that increased sucrose turnover in the vacuole and increased apoplastic uptake of reducing sugars. These findings extend our understanding of how some sugarcane varieties can naturally accumulate a high level of sucrose early in the season.     

相岭山系大、小熊猫主食竹类峨热竹的生长发育与环境因子间的相互关系

1994～1996年,在相岭山系冕宁县治勒自然保护区设点,对大熊猫和小熊猫主食竹类峨热竹与其它环境生态因子间的相互关系进行了长达3a的研究。研究结果表明,峨热竹的生长发育与环境生态因子如海拔、郁闭度、坡度和坡向之间有极其显著的相关关系,环境因子对其株高、基径、密度、发笋率、老笋比例和成竹死亡率等有较大的影响。随海拔高度的增加,竹子变矮变细,老笋比例下降;上层乔木郁闭度增加,竹子变稀,发笋率、老笋比例和成竹死亡率均降低。坡度增大,竹子变稀并且细而矮。阳坡的竹子率发笋高于阴坡。     

Cytology and ultrastructure of interactions between <Emphasis Type="Italic">Ustilago esculenta</Emphasis> and <Emphasis Type="Italic">Zizania latifolia</Emphasis>

Ustilago esculenta is a biotrophic smut fungus that parasitizes Zizania latifolia, an edible aquatic vegetable of the southern China region. Infection results in swelling of the upper parts of the Z. latifolia culm which are called jiaobai and have a unique flavor and delicacy and are popular among Chinese. The infection process of Z. latifolia by U. esculenta was investigated with light and electron microscopy. Distribution of hyphae was uneven in plants; hyphae were mainly present in the swollen upper parts (jiaobai), the nodal regions of mature culms and old rhizomes and buds or shoots. Hyphae were rare in the internodes of mature culms and were fewer in the internodes of old rhizomes. All new buds produced on the nodes of culms and rhizomes were infected by hyphae in November before and in March after overwintering. The hyphae grew into the buds from the parent nodes via intervascular tissues only or via parenchyma tissues and vascular bundles. Hyphae extended within and between the host cells and frequently formed hyphal aggregations or clusters, not only in the mature tissues but also in developing tissues. The typical interface between the fungal hyphal wall and invaginated host plasma membrane comprised a sheath. The sheath surrounding a hyphae comprised an outer electron-opaque matrix and an inner electron-dense layer. The electron-opaque matrix layers were thicker in jiaobai tissues, ranging from 0.28 to 0.85 μm. The electron-dense hyphal coatings were more conspicuous in the young buds or shoots and mature culms than in the jiaobai. The intercellular hyphae caused large cavity formation between the cells or rupture of host cell walls, for gaining entry into host cells. The broken host cell wall fused with the electron-opaque matrix of the hyphal sheath as an interactive interface. The teliospore wall and wall ornamentation development was the same in postmature jiaobai tissues with sporadic sori and in the huijiao (jiaobai tissues containing the massive sori), but a sheath enveloping the teliospore was more transparent in the process of teliospore development in the jiaobai than in the huijiao.     

Phragmites australis at an extreme altitude: Rhizome architecture and its modelling

In central EuropePhragmites australis is a lowland plant, occurring rarely up to the tree line. In the Velká Kotlina cirque (Jeseníky mountains, NE Czech Republic), where it reaches its maximum altitude at about 1350 m a.s.l., its culms are 0.5–0.7 m high and the plants flower only in some years. During the last decade no germinable seeds have been observed. The architecture ofPhragmites rhizomes from this site was studied on seven randomly selected clonal fragments. They consisted of 3 to 10 partial tussocks (clumps) and 4 to 17 green shoots. The total length of the rhizomes was 9.7 to 50 m per plant. The number of nodes per plant was 96 to 431 and the longest internodes were 83 mm long. The number of side branches was 31 to 105 per plant. The branching angle depended on the type of branched rhizome. The mean angles of horizontal rhizomes, which connect individual tussocks, were relatively wide (modus 45°, arithmetic mean 37°), whereas within a tussock much sharper angles of branching prevailed (modal value 5°, arithmetic mean 15°). The mean internode-to-internode angle on continuing rhizomes was about 8°, with a wide variation. An architectural, spatially-explicit model ofPhragmites rhizome growth has been developed, showing that thePhragmites population in the studied locality can be maintained by vegetative multiplication, and seedling recruitment is not needed for its long-term persistence.     

Impact of Culm Harvest on Seed Production in a Monocarpic Bamboo

Successful regeneration of bamboos from seed is a key issue in the ecology of many tropical regions and the livelihoods of their inhabitants. The gregarious monocarpy of many bamboos may be driven by a need to satiate seed predators by seeding in abundance at infrequent intervals. In long‐lived clonal monocarps, seed production is expected to be positively related to the success of the clone in generating more and larger ramets during its lifetime. Ramification may be constrained by harvesting of culms, but it is unclear whether the reduction in productivity is proportional to the loss of reproductive biomass. We counted the seed produced by 661 culms (ramet stems) sampled from 90 clumps of the gregariously monocarpic bamboo Schizostachyum dullooa that is intensively harvested by villagers in northeastern India. The smallest clumps had fewer culms and few or no culms more than one year old. Seed production was indeed positively related to culm size and the number of culms in a clump. First‐year culms were markedly more productive than older culms after controlling for culm diameter and clump size. There was a negative effect of clump size on productivity per culm which may occur because clumps that had been harvested heavily were able to exploit resources retained in rhizomes from harvested culms. Nevertheless, small clumps produced much less seed than larger clumps, generating a risk of unknown magnitude that heavily harvested stands of monocarpic bamboos may be unable to satiate seed predators during their single opportunity for reproduction.     

Papyrus swamp development in the Upemba Basin, Zaïre: studies of population structure in Cyperus papyrus stands

The ecology and distribution of papyrus swamps are briefly reviewed together with the ecology, distribution structure and growth characteristics of Cyperus papyrus.
Three age-classes of culm are recognized and girth and density of each are recorded along transects through mree swamps in the Upemba Basin of Zaïre. A regression of culm-unit biomass on culm girth, developed for Ugandan swamps, is used to convert Upemba measurements into biomasses. It is shown that simple measurements, taken at only a single point in time, yield much information on both the structure and the dynamics of papyrus populations.
It is suggested that the standing-crops of such populations are determined by available light, and by local temperature regimes, and are therefore constant, irrespective of site quality (nutrient status), for any particular climatic zone. Culm-duration (life-time) is not significantly influenced by climate and the ratio culms/rhizomes provides a useful indicator of comparative productivity. Unlike community biomass, productivity can be limited by site quality.     

Environmental determinants of Phragmites australis expansion in a New Jersey salt marsh: an experimental approach

Interdependence among disturbance events, ecosystem properties, and biological invasions often make causal relationships difficult to discern. For example, Phragmites australis invasion in mid-Atlantic salt marshes is often associated with disturbances that create well-drained features as well as with low sulfide concentrations, but explanations of these associations have been elusive. We tested experimentally: 1) that disturbances increasing wetland drainage facilitate Phragmites invasion by altering sulfide concentrations and salinity; 2) that translocation allows plants to spread beyond drainage areas; and 3) that plants can then lower edaphic stress through pressure ventilation of the rhizosphere and promote further expansion. At the invasion front, treatments of 1) severing rhizomes to halt translocation and 2) combined severing with clipping dead culms to limit ventilation of the rhizosphere killed most culms, but did not affect pore water chemistry. In already invaded areas, severing and clipping reduced culm height and panicle production, severing alone and in combination with clipping also raised sulfide and ammonium concentrations in the root zone. There were no treatment effects on plant performance or pore water chemistry along mosquito ditches, where sulfide concentrations were negligible. Small-scale hydrological alterations such as ditches appear to provide suitable sites for the establishment of Phragmites because soils are well-drained and are low in free sulfides. Subsequent expansion into more hostile areas occurs through translocation, with well-drained areas acting as sources for essential substances. Once established, the plant increases rhizosphere oxygenation and lowers sulfide concentrations.