Interactive effects of redox intensity and phosphate availability on growth and nutrient relations of Cladium jamaicense (Cyperaceae)

Expansion of Typha domingensis into areas previously dominated by Cladium jamaicense in the Florida Everglades has been linked to anthropogenic phosphorus (P) enrichment and increased hydroperiod. The principal stress factor for plants in flooded soils is biochemical reduction, the intensity of which is measured as redox potential (Eh). The objective of this study was to assess the growth response of C. jamaicense to Eh (-150, +150, and +600 mV) and P availability (10, 80, and 500 μg P/L). Plants were grown hydroponically in a factorial experiment using titanium (Ti(3+)) citrate as an Eh buffer. Treatment effects on growth, biomass partitioning, and tissue nutrients were recorded. Growth approximately doubled in response to a 50-fold increase in P availability. Low redox significantly reduced growth and tissue P concentration. While plant P concentrations increased 20-fold between the 10 and 500 μg P/L treatments, P concentrations were 50-100% higher at +600 mV than at -150 mV within each phosphate level. At high Eh, C. jamaicense appears well adapted to low nutrient environments because of its low P requirement and high retention of acquired P. However, at low Eh the ability to acquire or conserve acquired P decreases and as a consequence, higher phosphate levels are required to sustain growth. Findings of this study indicate that young C. jamaicense exhibits low tolerance to strongly reducing conditions when phosphate is scarce.     

Effects of salinity on growth and cation accumulation of Sporobolus virginicus (Poaceae)

Optimal growth of euhalophytes requires moderate concentrations of salt and, in dicotyledons, is associated with succulence and accumulation of Na(+) in plant tissues. However, reports of salt-stimulated growth in monocotyledons are rare. Relative growth rate (RGR), biomass accumulation, and water content were studied in Sporobolus virginicus (Poaceae), a C(4) chloridoid grass, grown hydroponically with different concentrations of NaCl. Cation concentrations were determined by atomic absorption spectrophotometry. Optimal growth occurred at 100-150 mmol/L NaCl and was not dependent on nitrogen levels or accompanied by accumulation of Na(+) in leaves. Biomass accumulation and RGR in plants grown at 450 mmol/L NaCl were greater than in plants grown at 5 mmol/L. The Na?:?K ratios were lower in leaves than in roots, indicating discrimination in Na(+) and K(+) transport. Secretion of Na(+) increased from 166.5 to 336.7 mmol · g(-1) dry biomass · d(-1) as the NaCl concentration of the nutrient solution increased from 125 mmol/L to 450 mmol/L. Water concentrations of leaves and shoots were significantly greater in plants grown at optimal levels of salinity than in plants grown at lower or higher salinities. These results demonstrate salt-stimulated growth in a monocotyledon.     

An architectural model for Eleocharis: morphology and development of Eleocharis cellulosa (Cyperaceae)

Species of Eleocharis are prominent in aquatic and wetland habitats and serve as models for study of physiological adaptations to aquatic environments. The genus has an unusual morphology because the major photosynthetic organ is the stem. In order to define an architectural model for the genus to understand the evolution of this morphology, we examined mature morphology and development of E. cellulosa in living and fixed material using light and scanning electron microscopy. Eleocharis cellulosa has sympodial, vertical shoots that produce the photosynthetic culms and horizontal shoots that mix monopodial and sympodial development. Each sympodial unit produces three bracts, an elongated photosynthetic internode, then a fourth bract and an inflorescence that either aborts on vegetative culms or expands on reproductive culms. On each sympodium, the first bract subtends a precocious axillary bud that reiterates the sympodial unit; the second bract subtends a bud that develops the horizontal shoot. In both horizontal and vertical shoots, the internode below the second bract is produced by both the second bract and the renewal shoot. Sympodial growth is present in seedlings. In other species of Eleocharis, the structure of the sympodial unit is conserved but morphological diversity develops from variation in horizontal shoot growth.     

The effect of an elevated atmospheric CO2 concentration on growth, photosynthesis and respiration of Plantago major

The effect of an elevated atmospheric CO₂ concentration on growth, photosynthesis and root respiration of Plantago major L. ssp. major L. was investigated. Plants were grown in a nutrient solution in growth chambers at 350 and 700 μl I⁻¹ CO₂ during 7 weeks. The total dry weight of the Co₂-enriched plants at the end of this period was 50% higher than that of control plants. However, the relative growth rate (RGR) was stimulated only during the first half of the growing period. The transient nature of the stimulation of the RGR was not likely to be due to end-product inhibition of photosynthesis. It is suggested that in P. major , a rosette plant, self-shading causes a decline in photosynthesis and results in an increase in the shoot: root ratio and a decrease in RGR. CO₂-enriched plants grow faster and cosequently suffer more from self-shading. Corrected for this ontogenetic drift, high CO₂ concentrations stimulated the RGR of P. major throughout the entire experiment.     

Compensatory Changes in the Partitioning of Dry Matter in Relation to Nitrogen Uptake and Optimal Variations in Growth

Equations are derived relating relative growth rate (RGR) toroot:shoot ratio, root length, nitrogen inflow rate, leaf area,photosynthesis and carbon and nitrogen concentrations in theplant. The extents to which changes in specific root lengthand root: shoot ratio can compensate for the effects of lowN availability upon RGR are examined. Such responses could haveseveral compensatory functions: maximizing RGR; maintaininggrowth in which the activities of root and shoot limit RGR equally;and maximizing the efficiency of increase in RGR. Growth, nitrogen, carbon, dry matter, partitioning, root:shoot ratio, relative growth rate     

Growth Rate, Photosynthesis and Respiration in Relation to Leaf Area Index

This work examined three possible explanations of growth rateresponses to leaf area index (LAI) in which growth rate perunit of ground area (crop growth rate, CGR) increased to a plateaurather than decreasing above an optimum LAI at which all lightwas intercepted. Single leaf photosynthetic measurements, andwhole plant 24 h photosynthesis and respiration measurementswere made for isolated plants and plants in stands using Amaranlhushybridus, Chenopodium album, and two cultivars of Glycine maxgrown at 500 and 1000 µimol m^–2 S^–1 photosyntheticphoton flux density at 25 °C. CGR, relative growth rate(RGR), and LAI were determined from 24 h carbon dioxide exchangeand leaf area and biomass measurements. Respiration increasedrelative to photosynthesis with crowding in A. hybridus andthere was an optimum LAI for CGR. In contrast, the ratio ofrespiration to photosynthesis was constant across plant arrangementin the other species and they had a plateau response of CGRto LAI. Neither increased leaf photosynthetic capacity at highLAI nor a large change in biomass compared to the change inLAI could account for the plateau responses. It was calculatedthat maintenance respiration per unit of biomass decreased withdecreasing RGR in C. album and G. max, but not A. hybridus,and accounted for the plateau response of CGR to LAI. Sincesimilar decreases in maintenance respiration per biomass atlow RGR have been reported for several other species, a constantratio of respiration to photosynthesis may occur in more speciesthan constant maintenance respiration per unit of biomass. Amaranlhus hybridus L., Chenopodium album L., Glycine max L Merr, soybean, photosynthesis, respiration, growth, leaf area index     

Nutrient and growth responses of cattail (Typha domingensis) to redox intensity and phosphate availability

Background and Aims

In the Florida Everglades, the expansion of cattail (Typha domingensis) into areas once dominated by sawgrass (Cladium jamaicense) has been attributed to altered hydrology and phosphorus (P) enrichment. The objective of this study was to quantify the interactive effects of P availability and soil redox potential (Eh) on the growth and nutrient responses of Typha, which may help to explain its expansion.

Methods

The study examined the growth and nutrient responses of Typha to the interactive effects of P availability (10, 80 and 500 µg P L⁻¹) and Eh level (−150, +150 and +600 mV). Plants were grown hydroponically in a factorial experiment using titanium (Ti³⁺) citrate as a redox buffer.

Key Results

Relative growth rate, elongation, root-supported tissue/root ratio, leaf length, lateral root length and biomass, as well as tissue nutrient concentrations, were all adversely affected by low Eh conditions. P availability compensated for the negative effect of low Eh for all these variables except that low P stimulated root length and nutrient use efficiency. The most growth-promoting treatment combination was 500 µg P L⁻¹/ + 600 mV.

Conclusions

These results, plus previous data on Cladium responses to P/Eh combinations, document that high P availability and low Eh should benefit Typha more than Cladium as the growth and tissue nutrients of the former species responded more to excess P, even under highly reduced conditions. Therefore, the interactive effects of P enrichment and Eh appear to be linked to the expansion of Typha in the Everglades Water Conservation Area 2A, where both low Eh and enhanced phosphate availability have co-occurred during recent decades.Key words: Everglades, growth, nutrient, phosphorus, redox potential, Typha domingensis     

Effects of root hypoxia and a low P supply on relative growth, carbon dioxide exchange rates and carbon partitioning in Pinus serotina seedlings

The present study was conducted to determine how 10 weeks of root hypoxia and a low P supply altered relative growth, and carbon acquisition and partitioning in a moderately flood tolerant pine. Pond pine (Pinus serotina Michx.) seedlings were grown in continuously flowing solution culture at 5 or 100 μM P, under aerobic or hypoxic solution conditions. Staggered harvests were used to ascertain changes in biomass allocation and relative growth over time. Carbon dioxide exchange rates (CER) were determined by infrared gas analysis, and needles were analyzed for inorganic P (Pi), sucrose, reducing sugars and starch. Although aeration treatment had no significant effect on shoot dry weight or shoot ontogeny, root dry weight of hypoxic seedlings was significantly lower than that of aerobic seedliings after 8 weeks, regardless of the P treatment. Mean relative growth rates (RGR) of roots in the high P treatment initially decreased under hypoxia, but recovered by the sixth week with the production of adventitious roots. Two weeks of hypoxic growth conditions decreased CER and stomatal conductances of seedlings in the high P treatment by more than 30% relative to their aerobic counterparts. Stomatal closure was not accompanied by a decrease in intracellular CO₂, but was accompanied by an increase in starch accumulation. Recoveries of CER, stomatal conductance and carbohydrate metabolism coincided with the recovery of root growth. Low P growth conditions did not significantly affect shoot or root dry weight until the sixth week of treatment. However, differences in seedling RGR, particularly needle RGR, were discernable during the first 2 weeks. Low P treatment effects on CER paralleled changes in needle RGR, with needle RGR more affected than CER. After 6 weeks, CER of aerobically grown seedlings in the low P treatment were only 15% lower than CER of seedlings in the high P treatment, despite a 31% and 75% reduction in needle RGR and Pi concentrations, respectively. Increased starch concentrations of recently expanded needles at this time were probably a result of diminished growth. The inhibitory effect of a low P supply on shoot growth, more specifically on needle expansion and emergence of new fascicular needles, probably limited net carbon fixed per plant more than any direct effect of low P on CER per se.     

Natural hybridization in tropical spikerushes of Eleocharis subgenus Limnochloa (Cyperaceae): Evidence from morphology and DNA markers

? Premise of the study: Natural hybridization represents an important force driving plant evolution and affecting community structure and functioning. Hybridization may be overlooked, however, among morphologically highly uniform congeners. An excellent example of such a group is Eleocharis subgenus Limnochloa, which has no reliably proven hybrids. Does this reflect biological barriers to interspecific crosses or difficulties in detecting the hybrids? We tested the hypothesis that hybridization occurs among sympatric Eleocharis cellulosa, E. interstincta, and E. mutata in northern Belize, Central America. ? Methods: Morphometric study (407 plants) was followed by examination of inter-simple sequence repeat (ISSR) polymorphisms (44 plants) and ITS sequence variation (33 plants). ? Key results: Two putatively hybrid morphotypes were discerned-E. cellulosa-resembling and E. interstincta-resembling. DNA markers of E. cellulosa and E. interstincta displayed additive constitution in plants from one E. cellulosa-resembling population only. The other putatively hybrid populations contained ISSR and ITS markers of the species they resembled morphologically, several unique ISSR markers, and ITS sequences of an undescribed South American Limnochloa entity. DNA markers of E. mutata were absent in the putative hybrids. ? Conclusions: Simultaneous use of various types of molecular markers can overcome many pitfalls of investigations concerning hybridization among closely related and morphologically similar species. Northern Belize represents a hybrid zone of E. cellulosa and E. interstincta. A third participant in the hybridization events occurring in this zone is an unknown Limnochloa lineage but is not E. mutata. Interspecific hybridization may play a significant role in the diversification of Eleocharis.     

赤霉素GA3调节黄芩组织培养中芽和根的形成

应用组织培养技术对黄芩进行外源激素调控研究。在培养不同时间进行的不同培养基之间的转移培养研究表明,组织培养条件下,在培养基中添加赤霉素,可显著刺激黄芩外植体芽的形成,同时抑制根的生长。在加有GA3的IAA培养基上,GA3显著影响黄芩组织培养物中的黄酮含量。在黑暗条件下,开始在2.5μmol/LIAA培养基中培养6d,随后转移到5μmol/LGA3培养基上培养,黄芩外植体中黄岑苷、黄岑素和汉黄芩苷的含量最高,分别为14.90,2.70和0.54μgmg-1(干重)。     

Waterlogging injury in sorghum seedlings and their kinetin effected rapid recovery

Short term waterlogging affected the growth of sorghum seedlings as indicated by a high mortality rate of the seedlings and a decrease in the shoot and root biomass, net assimilation rate (NAR), leaf area ratio (LAR), relative growth rate (RGR), total photosynthetic area and rate of photosynthesis. Plant resistance to the stress was exhibited by decreased stomatal index (SI), rate of transpiration and rate of respiration. When the stressed plants were exposed to normal conditions, the recovery of the seedlings was rapid and was enhanced by foliar application of kinetin at 5 mgl^-1.     

Differential responses of oat cultivars to phosphate deprivation: plant growth and acid phosphatase activities

The effect of phosphate starvation on growth and acid phosphatases (APases) localization and activity in oat tissues was investigated. Oat cultivars (Avena sativa L.??Arab, Polar, Szakal) were grown for 1?C3?weeks in complete nutrient medium (+P) and without phosphate (?P). Pi concentration in plant tissues decreased strongly after culturing on ?P medium. Pi deficit reduced shoot growth, stimulated root elongation and increased ratio of root/shoot in all oat cultivars. Pi deficit had a greater impact on growth of oat cv. Polar than other varieties. A decrease in the internal Pi status led to an increase of acid phosphatase activities in extracts from shoots and roots, and in root exudates. The highest activity of secreted APases was observed for oat cv. Arab, during the third week of growth under Pi-deficient conditions. The activity of extracellular APase was high in young, growing zones of roots of ?P plants. Histochemical visualization indicated high activity of APases in the epidermis and vascular tissues of ?P plants. Pi deficiency increased intracellular APase activity in shoot mainly in oat cv. Polar, whereas APase activity in roots was the highest in oat cv. Szakal. Protein extracts from roots and shoots were run on native discontinuous PAGE to determine which isoform(s) may be affected by Pi deficiency. Three major APase isoforms were detected in all oat plants; one was strongly induced by Pi deficit. The studied oat cultivars differed in terms of acclimation to deficiency of phosphate??used various pools of APases to acquire Pi from external or internal sources.     

SPECIES COMPOSITION,BIOMASS, AND NUTRIENT CONTENT OF PERIPHYTON IN THE FLORIDA EVERGLADES1

Periphyton biomass, nutrient dynamics in the biomass, and species composition were studied in two Florida Everglades sloughs from August 1991 to August 1992. Periphyton biomass on macrophytes was strongly season-dependent. Maximum biomasses, 1180, 161, and 59 g dry mass.m^?2 on Eleocharis vivipara, E. cellulosa, and Nymphaea odorata, respectively, occurred in summer and early autumn; winter and spring periphyton biomass was very low (practically not measurable). Periphyton was dominated by blue-green algae (cyanobacteria) during the summer and autumn; diatoms dominated during the winter and spring. Green algae occurred mostly during the summer and autumn, but their growth was sparse and did not contribute significantly to periphyton biomass. Nitrogen-to-phosphorus ratios in the periphyton were very high (59–121:1), suggesting phosphorus limitation of periphyton growth. The periphyton contained large concentrations of calcium (up to 22.3% on dry mass basis) especially in late summer and autumn.     

太平洋中西部海域浮游植物营养盐的潜在限制

2009年8月至9月期间在太平洋西部N1站和中部N2站进行现场营养盐加富培养实验。结果显示:N1站,浮游植物生物量对N或者P添加都有较强的响应,其中N+P+Si组和N+P组浮游植物长势迅速,叶绿素a从初始的0.03μg/L分别达到2.12μg/L和1.83μg/L,同时P先于N和Si之前被耗尽;说明N1站为N、P共同限制,P是首要限制因子。而N2站,浮游植物生物量仅对N、P共同添加有明显响应,N先于P和Si被浮游植物消耗殆尽。利用培养过程中营养盐比值变化推断,N1站浮游植物以低于Redfield ratio(16N∶1P)吸收N和P;而N2站浮游植物以高于Redfield ratio(16N∶1P)吸收N和P。这可能解释了太平洋西部的寡营养盐海域为潜在P限制,而在太平洋中部海域则为潜在N限制。     

Arsenic-resistant bacteria solubilized arsenic in the growth media and increased growth of arsenic hyperaccumulator Pteris vittata L

The role of arsenic-resistant bacteria (ARB) in arsenic solubilization from growth media and growth enhancement of arsenic-hyperaccumulator Pteris vittata L. was examined. Seven ARB (tolerant to 10 mM arsenate) were isolated from the P. vittata rhizosphere and identified by 16S rRNA sequencing as Pseudomonas sp., Comamonas sp. and Stenotrophomonas sp. During 7-d hydroponic experiments, these bacteria effectively solubilized arsenic from the growth media spiked with insoluble FeAsO? and AlAsO? minerals (from < 5 μg L?1 to 5.04-7.37 mg L?1 As) and enhanced plant arsenic uptake (from 18.1-21.9 to 35.3-236 mg kg?1 As in the fronds). Production of (1) pyochelin-type siderophores by ARB (fluorescent under ultraviolet illumination and characterized with thin layer chromatography) and (2) root exudate (dissolved organic C) by P. vittata may be responsible for As solubilization. Increase in P. vittata root biomass from 1.5-2.2 to 3.4-4.2 g/plant dw by ARB and by arsenic was associated with arsenic-induced plant P uptake. Arsenic resistant bacteria may have potential to enhance phytoremediation of arsenic-contaminated soils by P. vittata.     

Phosphorus–zinc interactions in cotton: consequences for biomass production and nutrient‐use efficiency in photosynthesis

The fragmentary information on phosphorus (P) × zinc (Zn) interactions in plants warrants further study, particularly in plants known for their high P and Zn requirements, such as cotton (Gossypium hirsutum L.). The objective of this study was to investigate the effect of P × Zn interactions in a modern cultivar of cotton grown hydroponically. Biomass, mineral nutrition and photosynthetic parameters were monitored in plants receiving contrasting combinations of P and Zn supply. Root biomass, length and surface area were similar in plants with low P and/or low Zn supply to those in plants grown with high P and high Zn supply, reflecting an increased root/shoot biomass quotient when plants lack sufficient P or Zn for growth. Increasing P supply and reducing Zn supply increased shoot P concentrations, whilst shoot Zn concentrations were influenced largely by Zn supply. A balanced P × Zn supply (4 mM P × 4 μM Zn) enabled greatest biomass accumulation, while an imbalanced supply of these nutrients led to Zn deficiency, P toxicity or Zn toxicity. Net photosynthetic rate, stomatal conductance, transpiration rate and instantaneous carboxylation efficiency increased as P or Zn supply increased. Although increasing P supply reduced the P‐use efficiency in photosynthesis (PUEP) and increasing Zn supply reduced the Zn‐use efficiency in photosynthesis (ZnUEP), increasing Zn supply at a given P supply increased PUEP and increasing P supply at a given Zn supply increased ZnUEP. These results suggest that agricultural management strategies should seek for balanced mineral nutrition to optimize yields and resource‐use efficiencies.     

Compensatory growth responses to clipping defoliation in Leymus chinensis (Poaceae) under nutrient addition and water deficiency conditions

Compensatory growth responses of Leymus chinensis, a dominant species in Inner Mongolia steppe, to clipping defoliation were evaluated in a pot-cultivated experiment under different nutrient (N and P) and water availability conditions. Leymus chinensis exhibited over-compensatory growth at the light and moderate clipping intensities (20% and 40% aerial mass removed) with a greater accumulated aboveground biomass, higher relative growth rate (RGR), more rhizomatic tillers and a stimulation of compensatory photosynthesis to the remnant leaves as compared with those of the unclipped plants. Intense clipping (80% aerial mass removed), which removed most of the aboveground tissues, greatly reduced the growth of aboveground biomass in comparison with that of the unclipped plants. Nitrogen addition only slightly improved the biomass production and RGR in light and moderately clipped plants, and it did not allow plants in the intense clipping condition to over-compensate. Phosphorus addition had no obvious influences on the growth and physiological responses to clipping defoliation. These results indicated that nutrient addition could not compensate for the negative effects of severe clipping on the defoliated grass. On the other hand, there were no distinct positive responses under water deficiency condition for L. chinensis at all clipping intensities with a significant reduction of aboveground and belowground biomass, lower RGR, fewer rhizomatic tillers, and a lower net photosynthetic rate than other wet treatments. Additionally, the chlorophyll contents of remnant leaves gradually increased with the increase of clipping intensities in each treatment. In conclusion, although L. chinensis could compensate for tissues removal by some morphological and physiological responses, intense clipping and drought can result in a significant decrease of biomass and growth rate, even under enriched nutrition conditions.     

Uptake of phosphorus and lead by Brassica juncea and Medicago sativa from chloropyromorphite

In situ remediation of lead (Pb)-contaminated soils via phosphate amendments has been extensively used to immobilize Pb as pyromorphite. However, in phosphorus (P) deficient soils, plants may develop extensive root systems to access P in any P-containing minerals, thereby affecting the stability of Pb5 (PO4)3Cl (Chloropyromorphite; CP). We grew Brassica juncea and Medicago sativa in sand culture to evaluate the stability of CP in the presence or absence of hydroxyapatite (HA) as P source. Treatments (per kilogram of sand) watered with P-nutrient solution were control [PC0, (without CP)], 1, and 5 g Pb as CP [PC1, and PC5] and 0.45 g P as HA (PA), and those of watered with P-free nutrient solution were 1 and 5 g Pb as CP [NC1 and NC5], 5 g Pb as CP plus 0.45 g P as HA [NAC5], and 0.45 g P as HA [NA]. Plants in NC1 and NC5 treatments showed stunted growth and reductions in shoot elongation and leaf size. Among CP treated pots, the highest shoot Pb uptake was observed in NAC5 treatment. The results suggested that Pb accumulation and translocation in the plants was markedly higher in P-sufficient conditions than in P-deficient conditions.     

Differences in developmental plasticity and growth rate among drought-resistant and susceptible cultivars of durum wheat (Triticum turgidum L. var. durum)

Understanding how growth and development of durum wheat cultivars respond to drought could provide a basis to develop crop improvement programmes in drought-affected tropical and subtropical countries. A greenhouse experiment was conducted to study the responses of five durum wheat cultivars to moisture stress at different developmental phases. Phenology, total dry matter (TDM), relative growth rate (RGR), leaf area ratio (LAR), net assimilation rate (NAR), leaf weight ratio (LWR), specific leaf area (SLA) and shoot:root ratio were compared. Pre-anthesis moisture stress delayed phenological development, whereas post-anthesis moisture stress accelerated it. TDM accumulation rate was different between drought-resistant and susceptible cultivars. RGR and its components changed with age and moisture availability. Drought-resistant cultivars had a high RGR in favourable periods of the growing season and a low RGR during moisture stress. In contrast, the drought-susceptible cultivar (Po) showed an opposite trend. LAR explained the differences in RGR (r=0.788) best, whereas the relationship between NAR and RGR was not significant. Even though both LWR and SLA were important factors determining the potential growth rate, LWR was of major importance to describe cultivar differences in LAR, and consequently in RGR. The drought-resistant cultivars Omrabi-5 and Boohai showed vigorous root development and/or a low shoot:root ratio. It is concluded that biomass allocation is the major factor explaining variation in RGR among the investigated durum wheat cultivars.     

几种营养物质对烟草细胞生长和CoQ10含量的影响

研究了蔗糖、KH₂PO₄、NH₄NO₃比对烟草细胞生长和CoQ₁₀量的影响,结果表明,当蔗糖浓度为30g/L时,CoQ₁₀总量最高,此时细胞产量、CoQ₁₀含量和总量分别为19.8g/L、414.7μg/g(dwt)、8212μg/L。在上述蔗糖浓度下,当KH₂PO₄起始浓度为340mg/L、NH₄NO₃为12时,细胞产量、CoQ₁₀含量和总量分别最高,高于此比例时有利于CoQ₁₀形成 ,但不利于细胞生长。