Characterization of suspended solids in Lake Biwa by measuring their elemental composition of Al,Si, P,S, K,Ca, Ti,Mn, and Fe

We investigated the concentrations of particulate Al, Si, P, S, K, Ca, Ti, Mn, and Fe at various areas of Lake Biwa, such as the limnetic zone in the northern basin, the offshore zone, the dredged area and Akanoi Bay in the southern basin, and reed community areas in the shore. In the wide and deep northern basin, the density of suspended solids was low, and substantial fractions of particulate Si, P, S, and Ca were biogenic matter produced by phytoplankton. In the offshore zone in the narrow and shallow southern basin, the concentration ratios of particulate Si, K, Ca, Ti, and Fe against particulate Al were almost constant throughout the year and were similar to those found in crusts. Therefore, these elements are most likely derived from terrigenous clay matter. In the dredged area where the hypolimnion became anoxic during the stratification period, the concentrations of particulate Fe, Mn, P, S, and Ca increased because of the formation of hydrous Fe and Mn oxides and Fe sulfide. In Akanoi Bay, the ratios of Mn/Al and Fe/Al were high because of the stirring of sediments enriched with Mn and Fe oxides. In the reed community areas, the concentration of suspended solids correlated well with particulate Fe and P, but not with Al or chlorophyll a; therefore these areas seem to be rich in humic organic matter enriched with P and Fe due to microbial activity. Thus, the elemental composition of suspended solids reflects the chemical, biological, and physical processes in the lake and gives us useful indices of the characteristics of the water. Received: February 5, 2001 / Accepted: August 28, 2001     

Chemical composition of modern and pre-acidification sediments in the Tatra Mountain lakes

Concentrations of major nutrients (C, N, P) and acid soluble metals (Ca, Mg, K, Al, Fe, Mn, Pb, and Zn) were determined in modern (0–1 cm) and pre-acidification (5–10 cm) sediment layers collected from 37 alpine and 3 forest lakes in the Tatra Mountains (Slovakia, Poland) in 1996–1998. Sediment composition reflected catchment characteristics and productivity of lakes. In the sediments of alpine lakes, C and N concentrations decreased and Mg increased with a decreasing proportion of vegetation and soil in the catchment. Decreasing Ca:Mg ratios in sediments along the vegetation gradient was inverse to that in water, and could be associated with different ratios of cations in water leachate from catchments and in solids which enter the lake due to soil erosion. Phosphorus concentrations increased with the proportion of moraine areas, with till soils rich in P. Concentrations of C, N, P, and Ca in sediments positively correlated to their concentrations in water. Sediment concentrations of Al and Al:Ca ratios increased with decreasing sediment and water pH. A negative correlation between water pH and concentrations of organic C in water and sediments indicated the important impact of organic acids on the acid status of the lakes exposed to higher terrestrial export of organic matter. Compared to the pre-acidification period, the modern sediments had significantly higher Fe, Mn, Zn, Pb, and K, but lower Mg concentrations. The Zn and Pb enrichment was more evident in oligotrophic alpine lakes than in more productive forest lakes and was independent of lake water or sediment pH. Fe and Mn concentrations in the modern sediments were higher than in ambient soils and bedrock, while those in pre-acidification sediments were similar to contemporary soils and to the rock layer. The enrichment of the modern sediments with Fe and Mn thus probably resulted from both their redox recycling and ecosystem acidification.     

Distribution of N,P, K,Ca, Mg,S, Zn,Fe, Mn and Cu in groundnut

Summary Concentration of N, P, K, Ca, Mg and S in summer groundnut crop was higher than in kharif while Zn, Fe, Mn and Cu contents were higher in summer crop. Kernel's N, P and Zn; Leaflet's Ca and Mn; Stem's K and Fe; Root's S and Cu and Petiole's Mg contents were highest. Shell's N, P, K, Mg, S, Zn and Cu; Kernel's Ca, Fe and Mn contents were the least. N, P, K, S, Zn and Cu concentrations decreased linearly as the crop grew. Ca, Mg, Fe and Mn concentrations did not display any distinct pattern. Ca concentration was positively correlated with pod yield in both the seasons.     

不同竹龄雷竹中硅及其他营养元素吸收和积累特征

在浙江省临安市的雷竹主产区,分别采集不同竹龄(1～4 a)和不同器官(叶、枝、秆)的雷竹样品,分析了Si和其他营养元素含量、吸收和积累特征,以及Si和其他营养元素之间的相互关系.结果表明： 雷竹各器官中C含量的大小顺序为竹秆>竹枝>竹叶,Si、N、P、K、Ca、Mg、Al、Fe和Mn含量的大小顺序为竹叶>竹枝>竹秆.除Mn主要积累在竹叶中外,其他9种营养元素主要积累在1年生雷竹的秆中.3～4年生雷竹竹叶的Si平均含量为13.66 g · kg^-1. 雷竹属于Si积累植物.随竹龄的增加,雷竹叶中的N、P、K和Mg含量减少,C、Al和Mn含量增加.雷竹对Si的吸收主要集中在第2年(57.1%),对N和K的吸收主要集中在前两年(67.7%～93.7%),此后N和K从植株体内流出,其流失量分别占总积累量的19.1%～39.1%.雷竹中Si与Ca、Al、Mn呈显著正相关,与N、P、K、Mg呈显著负相关.     

Effects of nitrogen enrichment on zooplankton biomass and N:P recycling ratios across a DOC gradient in northern-latitude lakes

We used data from whole-lake studies to assess how changes in food quantity (phytoplankton biomass) and quality (phytoplankton community composition, seston C:P and N:P) with N fertilization affect zooplankton biomass, community composition and C:N:P stoichiometry, and their N:P recycling ratio along a gradient in lake DOC concentrations. We found that despite major differences in phytoplankton biomass with DOC (unimodal distributions, especially with N fertilization), no major differences in zooplankton biomass were detectable. Instead, phytoplankton to zooplankton biomass ratios were high, especially at intermediate DOC and after N fertilization, implying low trophic transfer efficiencies. An explanation for the observed low phytoplankton resource use, and biomass responses in zooplankton, was dominance of colony forming chlorophytes of reduced edibility at intermediate lake DOC, combined with reduced phytoplankton mineral quality (enhanced seston N:P) with N fertilization. N fertilization, however, increased zooplankton N:P recycling ratios, with largest impact at low DOC where phytoplankton benefitted from light sufficiently to cause enhanced seston N:P. Our results suggest that although N enrichment and increased phytoplankton biomass do not necessarily increase zooplankton biomass, bottom-up effects may still impact zooplankton and their N:P recycling ratio through promotion of phytoplankton species of low edibility and altered mineral quality.

     

Soil-plant element relationships in a tundra ecosystem

The objectives of this study were to (1) provide a baseline estimate of soil and plant element concentrations for an intensive research site at Imnavait Creek in northern Alaska, and (2) examine the relationships between soil and plant elements in an arctic ecosystem. Soil and plant element concentrations were highly variable along biotic, spatial, and temporal axes. Deciduous shrubs had higher leaf concentration of N, P. K and Mg, whereas an evergreen shrub had higher leaf concentrations of Ca, Mn, Al and Si. Based on high required solution phase turnover rates, the most likely elemental deficiencies are N > P > K > Ca = Mg. Based on low required solution phase turnover rates and high soil concentrations. Fe, Mn, Zn and Cu deficiencies are unlikely. Manganese could be present in toxic concentrations. The nutrient bottleneck in tundra ecosystems appears to be the rate of nutrient movement to the solution phase.     

Fast and Inexpensive Detection of Total and Extractable Element Concentrations in Aquatic Sediments Using Near-Infrared Reflectance Spectroscopy (NIRS)

Adequate biogeochemical characterization and monitoring of aquatic ecosystems, both for scientific purposes and for water management, pose high demands on spatial and temporal replication of chemical analyses. Near-infrared reflectance spectroscopy (NIRS) may offer a rapid, low-cost and reproducible alternative to standard analytical sample processing (digestion or extraction) and measuring techniques used for the chemical characterization of aquatic sediments. We analyzed a total of 191 sediment samples for total and NaCl-extractable concentrations of Al, Ca, Fe, K, Mg, Mn, N, Na, P, S, Si, and Zn as well as oxalate- extractable concentrations of Al, Fe, Mn and P. Based on the NIR spectral data and the reference values, calibration models for the prediction of element concentrations in unknown samples were developed and tested with an external validation procedure. Except Mn, all prediction models of total element concentrations were found to be acceptable to excellent (ratio of performance deviation: RPD 1.8–3.1). For extractable element fractions, viable model precision could be achieved for NaCl-extractable Ca, K, Mg, NH₄ ⁺-N, S and Si (RPD 1.7–2.2) and oxalate-extractable Al, Fe and P (RPD 1.9–2.3). For those elements that showed maximum total values below 3 g kg⁻¹ prediction models were found to become increasingly critical (RPD <2.0). Low concentrations also limited the performance of NIRS calibrations for extracted elements, with critical concentration thresholds <0.1 g kg⁻¹ and 3.3 g kg⁻¹ for NaCl and oxalate extractions, respectively. Thus, reliable NIRS measurements of trace metals are restricted to sediments with high metal content. Nevertheless, we demonstrated the suitability of NIRS measurements to determine a large array of chemical properties of aquatic sediments. The results indicate great potential of this fast technique as an analytical tool to better understand the large spatial and temporal variation of sediment characteristics in an economically viable way.     

Direct and Indirect Effects of Dissolved Organic Matter Source and Concentration on Denitrification in Northern Florida Rivers

Using a natural gradient of dissolved organic carbon (DOC) source and concentration in rivers of northern Florida, we investigated how terrestrially-derived DOC affects denitrification rates in river sediments. Specifically, we examined if the higher concentrations of DOC in blackwater rivers stimulate denitrification, or whether such terrestrially-derived DOC supports lower denitrification rates because (1) it is less labile than DOC from aquatic primary production; whether (2) terrestrial DOC directly inhibits denitrification via biochemical mechanisms; and/or whether (3) terrestrial DOC indirectly inhibits denitrification via reduced light availability to—and thus DOC exudation by—aquatic primary producers. We differentiated among these mechanisms using laboratory denitrification assays that subjected river sediments to factorial amendments of NO₃ ^? and dextrose, humic acid dosing, and cross-incubations of sediments and water from different river sources. DOC from terrestrial sources neither depressed nor stimulated denitrification rates, indicating low lability of this DOC but no direct inhibition; humic acid additions similarly did not affect denitrification rates. However, responses to addition of labile C increased with long-term average DOC concentration, which supports the hypothesis that terrestrial DOC indirectly inhibits denitrification via decreased autochthonous production. Observed and future changes in DOC concentration may therefore reduce the ability of inland waterways to remove reactive nitrogen.     

Influence of <Emphasis Type="Italic">Potamogeton crispus</Emphasis> growth on nutrients in the sediment and water of Lake Tangxunhu

An incubation experiment was performed on Potamogeton crispus (P. crispus) using sediment collected from Lake Tangxunhu in the center of China, in order to determine the effects of plant growth on Fe, Si, Cu, Zn, Mn, Mg, P, and Ca concentrations in the sediments and overlying waters. After 3 months of incubation, Ca, Mg, and Si concentrations in the water column were significantly lower, and P and Cu concentrations were significantly higher than in unplanted controls. The effect of P. crispus growth on sediment pore waters and water-extractable elements varied. Concentrations of Ca, Mg, Si, Fe, Cu, and Zn were significantly higher, and P was significantly lower, than in pore waters of the control. Water-extracted concentrations of Fe, Mg, and Si in the sediments were lower, and P was higher, than in the control. Presence of P. crispus generally enhanced concentration gradients of elements between pore waters and overlying waters but not for P. The growth of P. crispus was associated with an increase in water pH and formation of root plaques, resulting in complex effects on the sediment nutritional status. Handling editor: S. M. Thomaz     

The relative importance of top-down and bottom-up control of phytoplankton in a shallow macrophyte-dominated lake

1. Mechanisms stabilizing the plant-dominated clear-water state were investigated in Little Mere, U.K. Replicated, factorial, mesocosm experiments, carried out in 1995 and 1996, were designed to investigate the relative importance of top-down (zooplankton grazing) and bottom-up (nitrogen-limitation) control in limiting algal growth, and the role of macrophytes in these processes. Treatments included increased salinity (1995) and sticklebacks (1996) to reduce zooplankton numbers, weekly nitrate additions and removal of macrophytes. 2. Contrary to the results of other studies, submerged plants did not reduce nitrate concentrations. Owing to the high stickleback density in the enclosures with fish, macrophytes did not provide a refuge for zooplankton during the experiment. In Little Mere, however, where fish densities are lower, macrophytes probably play a key role in maintaining clear water by providing refuge for pelagic zooplankton and habitat for attached Cladocera. 3. Phytoplankton in Little Mere was not nitrogen- (N) limited during the growing season. Although nitrogen availability sets a maximum potential phytoplankton biomass it was not realized owing to control by zooplankton grazing.     

Limnology and trophic status of glacial lakes in the tropical Andes (Cajas National Park,Ecuador)

相似文献   

Macrophyte induced microchemical changes in the water column of a northern Boreal Lake

Water quality was compared between open water and vegetated regions of the littoral zone of a Boreal lake. Within region variation occurred in vegetated areas and was species dependent. In the water lily versus the open water stations, conductivity and concentrations of B, Ca, Fe, K, Mg, Mn, S and Sr were elevated in the water column while N concentrations were lower. Wild rice areas were characterized by lower S and higher conductivity and Ca and Fe concentrations than open water areas. Variations in water quality in the vegetated regions occurred as a result of chemical exchange with the sediment in the proximity of the vegetation. Elemental concentrations in the sediment appeared to vary as a result of root aeration and nutrient uptake by the plants. Open water stations had elevated sediment levels of N, P and Al while elevated levels of Na were present in both the open water and wild rice plots. Water lily stations exhibited higher pH levels and higher concentrations of Fe, Mn and Ca than the open water plots. Plant tissue analysis indicated that between species elemental variation existed as well. Water lily tissue had higher concentrations of N, P, Mn, Zn, Ca, K and Mg than that of wild rice. It was postulated that aquatic macrophytes can influence the redox level of sediments and thereby dramatically alter the overlying water column forming microchemical environments in stagnant regions of the littoral zone.     

Hydrologic and biogeochemical controls on trace element export from northern Wisconsin wetlands

Wetlands play an important role in determining the water quality of streams and are generally considered to act as a sink for many reactive species. However, retention of chemical constituents varies seasonally and is affected by hydrologic and biogeochemical processes including water source, mineral weathering, DOC and SPM cycling, redox status, precipitation/dissolution/adsorption, and seasonal events. Relatively little is known about the influence of these factors on trace element cycling in wetland-influenced streams. To explore the role of wetlands with respect to the retention/release of trace elements to streams, we examined temporal and spatial patterns of concentrations of a large suite of trace elements (via ICP-MS) and geochemical drivers in five streams and wetland rivulets draining natural wetlands in a northern Wisconsin watershed as well as in their groundwater sources (terrestrial recharge, lake recharge, and older lake recharge). We performed principal components analyses of the concentrations of elements and their geochemical drivers in both the streams and rivulets to assist in the identification of factors regulating trace element concentrations. Variation in trace and major element concentrations among the streams was strongly related to the proportion of terrestrial recharge contributing to the stream. A dominant influence of water source on rivulet chemistry was supported by association of groundwater-sourced elements (Ba, Ca, Cs, Mg, Na, Si, Sr) with the primary statistical factor. DOC appeared in the first principal component factor for the streams and in the second factor for the rivulets. Strong correlations of Al, Cd, Ce, Cu, La, Pb, Ti, and Zn with DOC supported the important influence of DOC on trace metal cycling. A number of elements in the rivulets (Al, La, Pb, Ti) and streams (Al, Ce, Cr, Cu, La, Pb, Ti, Zn) had a significant particulate cycle. Redox cycling and precipitation/dissolution reactions involving Fe and Mn likely impacted Cu and Mo as evidenced by the low levels in the rivulets. Variance in Fe, Mn and the metal oxy-anions was associated with factors related to redox cycling and adsorption reactions in the wetland sediments. In streams, DOC and metals with a high affinity for DOC were associated with a factor which also included negative loadings for groundwater-sourced elements, reflecting the importance of seasonal hydrologic events which flush DOC and metals from wetland sediments and dilute groundwater sourced metals. Redox processes were of secondary importance in the streams but of primary significance in the rivulets, documenting the importance of anoxic conditions in wetland sediments on groundwater en route to the stream.     

Apparent and Measured Rates of Nitrification in the Hypolimnion of a Mesotrophic Lake

Three distinct phases were observed in the change of dissolved inorganic nitrogen concentrations in the hypolimnion of Grasmere. The second phase of decreasing ammonia and increasing nitrate concentrations was typical of the nitrification process. Observations on nitrate concentration gradients between surface sediments and the water column and experiments using the nitrification inhibitor N-Serve indicated the in situ activity of chemolithotrophic nitrifying organisms. Nitrification rates were estimated throughout the period of stratification by using the N-Serve and [¹⁴C]bicarbonate uptake method. Comparison of the field nitrate concentrations with the predicted nitrate concentrations (from estimates of the nitrification rate) indicated that the method underestimated the true rate of nitrification. Possible reasons for this are discussed.     

Lake Kinneret dissolved oxygen regime reflects long term changes in ecosystem functioning

Lake Kinneret (Israel) has undergone several prominent chemical and biological changes since 1970. Between 1970 and 1991 significant, long-term gradual increase were recorded in epilimnetic dissolved oxygen (DO) concentrations (about 20%), and in pH levels (0.2 units). Concomitantly there was a significant increase in hypolimnetic H₂S concentrations (about 75%) and a long-term gradual drop in zooplankton biomass (50%). Since 1994 these trends were reversed and the levels of the three chemical parameters have returned to those found in the 1970's and that of zooplankton to mid 1980's levels. The present study is an attempt to relate some of these long term changes by means of yearly oxygen budgets, based on fluxes of oxygen producing and consuming processes. This analysis raises the possibility that part of the long-term increase in epilimnetic DO and pH between 1970 to 1990 may be attributed to reduced inputs of organic matter from alochthonous sources and possibly to enhanced burial of organic matter in the bed sediments. However, the major cause for the observed increase in epilimnetic DO and pH is increased sedimentation of organic matter to the hypolimnion during stratification. As indicated by the amount of H₂S formed in the hypolimnion during stratification added to the amount of oxygen entrapped in this layer at the onset of thermal stratification, between 1970 to 1991 the sedimentation flux of organic matter increased by approximately 40%. It is estimated that during these two decades hypolimnetic respiration increased from ca. 8% of the annual amount of oxygen evolved due to photosynthesis during the early 1970's to ca. 12.5% during the 1980's. The shift in the layer of oxidative processes is suggested to be the result of a multi-annual decline in zooplankton grazing pressure, which led to increased sedimentation of organic matter. The reversed trends for DO, pH and H₂S since 1994 may have partially been due to the increase in zooplankton activity and partially due to changes in phytoplankton community structure.     

亚热带湿地松叶片多元素化学计量与养分回收对氮添加的短期响应

在我国南方亚热带湿地松人工林设置了3个水平的野外氮添加控制试验(0、40、120 kg N·hm^-2·a^-1),于2014和2015年生长季高峰期(7月底)和末期(10月底)采集湿地松成熟绿叶和落叶,分析外源氮添加对湿地松叶片碳(C)、氮(N)、磷(P)、钾(K)、钙(Ca)、镁(Mg)、铝(Al)、铁(Fe)、锰(Mn)9种元素浓度及其养分回收的影响.结果表明: N添加显著增加了湿地松绿叶中N、Al、Mn浓度,降低了P和2014年的Ca浓度,而对C、K、Mg、Fe 浓度无显著影响.N添加显著提高了绿叶N/P,且该比值及绿叶养分浓度(N、P、Mn)对N添加的响应依赖于N的剂量(高N条件下响应更强).N添加显著降低了2015年N的回收效率,提高了2014年K的回收效率.相比于养分回收效率,回收能力对增加的可利用氮响应更强.N添加显著降低了N的回收能力,提高了P、K的回收能力,降低了枯叶中的Fe浓度,而对枯叶中Ca、Mg、Al、Mn浓度无显著影响.这表明,N添加对叶片化学计量的影响因不同元素而异,植物会通过调整自身的养分内循环(养分回收)来应对环境变化.N添加提高了绿叶N/P和K/P,说明氮添加条件下植物生长可能由N、P共同限制转变为P限制.氮添加增加了绿叶中Al、Mn浓度,表明N添加下湿地松面临潜在的金属离子毒性风险升高.     

黔中喀斯特山区村域稻田土壤理化特性的空间变异特征及空间自相关性

以贵州省安顺市西秀区旧州镇文星村为研究区域,采用地统计学方法,研究了黔中喀斯特山区土壤容重、总孔隙度、毛管持水量、毛管孔隙度、通气孔隙度等物理特性与土壤pH、有机质、速效养分(S、Si、N、P、K、Fe、Zn、Cu、Mn)、全量养分(N、P、K)、交换性盐(Ca、Mg)及阳离子交换量等化学特性的空间变异特征及自相关性。在土壤各特征值中,pH、毛管孔隙度变异系数分别为4.59%、8.28%、9.83%,为弱变异;其他变异系数在10%-100%之间,为中等程度变异。半方差分析表明,土壤毛管持水量、通气孔隙度、毛管孔隙度、速效养分(S、Si、Cu、Mn)、全量养分(N、P、K)、交换性盐(Ca、Mg)及阳离子交换量的C₀/(C₀+C)<25%,表现为强烈的空间自相关性;容重、pH、速效养分(N、P、K、Fe、Zn)及有机质的C₀/(C₀+C)在25%-75%之间,表现为中等空间自相关性,而Moran''s I指数分析表明,除毛管孔隙度、有效S、速效P及有效Mn的空间自相关性较弱外,其他均呈正的显著空间自相关性。根据克里金插值图,在研究区域内土壤毛管孔隙度及毛管持水量与pH、速效养分(S、Si、Zn、Cu、)全P、全N、有机质、交换性Ca、Mg及阳离子交换量在东北方向和西北角分布较低,在西南角方向土壤容重及通气孔隙度分布较低,而pH、速效K、有效Cu、全P、全K、有机质、阳离子交换量分布较高,且速效养分(N、K、S、Cu)、全P、有机质的含量及阳离子交换量均存在从西南向东北方向递减的趋势。     

Experimental measurement of sediment nitrification and denitrification in Hamilton Harbour,Canada

This research examines the role of sediment nitrification and denitrification in the nitrogen cycle of Hamilton Harbour. The Harbour is subject to large ammonia and carbon loadings from a waste-water treatment plant and from steel industries. Spring ammonia concentrations rapidly decrease from 4.5 to 0.5 mg 1⁻¹, while spring nitrate concentrations increase from 1 to 2 mg l⁻¹, by mid-summer. A three-layer sediment model was developed. The first layer is aerobic; in it, oxidation of organics and nitrification occurs. The second layer is for denitrification, and the third layer is for anaerobic processes. Ammonia sources for nitrification include diffusion from the water column, sources associated with the oxidation of organics, sources from denitrification and from anaerobic processes. Diffusion of oxygen, ammonia and nitrate across the sediment-water interface occurs. Temperature effects are modelled using the Arrhenius concept. A combination of zero-order kinetics for nitrate or ammonia consumption with diffusion results in a half-order reaction, with respect to the water column loss rate to sediments. From experimental measurement, the rate of nitrification is 200 mg N 1⁻¹ sediment per day, while that of denitrification is 85 mg N 1–1 sediment per day at 20 °C. The Arrhenius activation energy is estimated as 15 000 cal/ mole-K and 17 000 cal/ mole-K for nitrification and denitrification, respectively, between 10 °C and 20 °C. Calculations of the flux of ammonia with the sediments, using the biofilm model, compare favourably with experimental observations. The ammonia flux from the water column is estimated to account for 20% of the observed decrease in water column stocks of ammonia, while the nitrate flux from the water column is estimated to account for 25% of the total nitrogen produced by the sediments.     

MINERAL CONCENTRATIONS IN AN AUTOPARASITIC PHORADENDRON CALIFORNICUM GROWING ON A PARASITIC P. CALIFORNICUM AND ITS HOST,CERCIDIUM FLORIDUM

Tissue concentrations of 12 mineral elements (Al, Ca, Cu, Fe, K, Mg, Mn, amino-N, Na, P, Si, and Zn) were measured in an autoparasitic mistletoe (Phoradendron californicum), the parasitic mistletoe on which it was growing (Phoradendron californicum), and the host tree (Cercidium floridum). Mineral concentrations in the autoparasite were typically 1.1–1.3 times higher than in the parasite. Mineral concentrations of all elements except Ca, Fe, Mg, Mn, and Si were higher in the parasitic mistletoe than its host. Mineral concentration differences are discussed relative to accumulation via the transpiration stream and translocation within the host via the phloem.