Genotype Variability in Photosynthetic Characteristics in Finger Millet

High variability in leaf gas exchange and related traits were found in 30 genotypes of field grown finger millet. The variability in carbon exchange rate per unit leaf area (P _N) can be partly attributed to the differences in the stomatal conductance (g_s) and area leaf mass (ALM). The P _N was positively correlated with total dry matter (TDM). However, no relationship between P _N and seed yield was found. The leaf area showed a positive and significant correlation with total biomass. None of the other gas-exchange traits had significant relationship either with TDM or with seed yield. The ALM showed a strong positive association with P _N. However, it was not correlated with either total biomass or seed yield. As a result, the use of ALM as surrogate for P _N for identifying high biomass producing genotypes only had a limited value. Hence selection for high P _N would result in higher biomass producing types.     

Variations in photosynthetic rate and associated parameters with age of oil palm leaves under irrigation

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in an adult oil palm (Elaeis guineensis) canopy were highest in the 9^th leaf and progressively declined with leaf age. Larger leaf area (LA) and leaf dry mass (LDM) were recorded in middle leaves. P _N showed a significant positive correlation with g _s and a negative relationship with leaf mass per area (ALM). The oil palm leaf remains photosynthetically active for a longer time in the canopy which contributes significantly to larger dry matter production in general and greater fresh fruit bunch yields in particular.     

Cotton genotypic variation in the photosynthetic response to irradiance

The photosynthetic response of 8 cotton (Gossypium hirsutum L.) genotypes to changing irradiance was investigated under field conditions during the 1998 through 2000 growing seasons. Equations developed to describe the response of net photosynthetic rate (P_N) to photosynthetic photon flux density (PPFD) demonstrated that, across all irradiances, the two okra leaf-type genotypes photosynthesized at a greater rate per unit leaf area than all of the six normal leaf-type genotypes. This superior photosynthetic performance of the okra leaf-type genotypes can be partially explained by their 13 % greater leaf chlorophyll content relative to that of the normal leaf-type genotypes. The 37 % reduction in leaf size brought upon by the okra leaf trait may have concentrated the amount of photosynthetic machinery per unit leaf area. Nevertheless, the lack of sufficient canopy leaf surface area suppressed the potential yield development that could accompany the higher P_N per unit leaf area.This revised version was published online in March 2005 with corrections to the page numbers.     

Influence of Potassium Deficiency on Photosynthesis,Chlorophyll Content,and Chloroplast Ultrastructure of Cotton Plants

In cotton (Gossypium hirsutum L.) grown in controlled-environment growth chamber the effects of K deficiency during floral bud development on leaf photosynthesis, contents of chlorophyll (Chl) and nonstructural saccharides, leaf anatomy, chloroplast ultrastructure, and plant dry matter accumulation were studied. After cotton plants received 35-d K-free nutrient solution at the early square stage, net photosynthetic rate (P _N) of the uppermost fully expanded main-stem leaves was only 23 % of the control plants receiving a full K supply. Decreased leaf P _N of K-deficient cotton was mainly associated with dramatically low Chl content, poor chloroplast ultrastructure, and restricted saccharide translocation, rather than limited stomata conductance in K-deficient leaves. Accumulation of sucrose in leaves of K-deficient plants might be associated with reduced entry of sucrose into the transport pool or decreased phloem loading. K deficiency during squaring also dramatically reduced leaf area and dry matter accumulation, and affected assimilate partitioning among plant tissues.     

Assessment of Genotypic Variation in Salt Tolerance of early CIMMYT Hexaploid Wheat Germplasm Using Photosynthetic Capacity and Water Relations as Selection Criteria

Twenty-five genotypes of early CIMMYT hexaploid wheat were screened for salt tolerance in a glasshouse experiment using photosynthetic capacity and water relation parameters as selection criteria. Under salt stress (150 mM NaCl) the genotypes Frontana, Norin-10, Mayo-54, Noreste-66, and Yaktana-54 excelled all other lines in shoot dry mass, and Na(20)TPP, Penjamo-62, Inia-66, Frontana, Siete Cerros, and Jaral-66 in grain yield per plant in both absolute and relative (percent of control) terms. Although net photosynthetic rate (P _N) declined in all genotypes due to salt stress, it was not helpful in discriminating among genotypes according to salt tolerance. Similarly, no positive relationships of salt tolerance of the genotypes with stomatal conductance, leaf water potential, or turgor pressure were found. Every genotype used its own specific mechanism to tolerate salt stress. However, a large amount of variation in salt tolerance observed in 25 early CIMMYT wheat genotypes can be of considerable practical value for improving salt tolerance in the existing commercial hexaploid wheats. This revised version was published online in August 2006 with corrections to the Cover Date.     

Comparison of Whole System of Carbon and Nitrogen Accumulation Between Two Maize Hybrids Differing in Leaf Senescence

A field experiment was conducted to investigate the carbon (C) and nitrogen (N) balance in relation to grain formation and leaf senescence in two different senescent types of maize (Zea mays L.), one stay-green (cv. P3845) and one earlier senescent (cv. Hokkou 55). In comparison with Hokkou 55, P3845 had a higher N concentration (N_c) in the leaves and a higher specific N absorption rate by roots (SAR_N), which indicated that a large amount of N was supplied to the leaves from the roots during maturation. This resulted in a higher photosynthetic rate, which supports saccharide distribution to roots. Thus, stay-green plants maintained a more balanced C and N metabolism between shoots and roots. Moreover, the coefficients of the relationship between the relative growth rate (RGR) and N_c, and between the photon-saturated photo-synthetic rate (P _sat) and N_c were lower in P3845. The P _sat per unit N_c in leaves was lower in the stay-green cultivars, which indicated that high yield was attained by longer green area duration and not by a high P _sat per unit N_c in the leaf. Consequently, a high P_sat caused a high leaf senescence rate because C and N compounds will translocate actively from the leaves.     

Activity of 1-Aminocyclopropane Carboxylic Acid Synthase in Two Mustard (Brassica juncea L.) Cultivars Differing in Photosynthetic Capacity

The pattern of activity of 1-aminocyclopropane carboxylic acid synthase (ACS) was similar to photosynthetic and growth traits observed at 30, 45, and 60 d after sowing in mustard (Brassica juncea L.) cultivars Varuna and RH 30 differing in photosynthetic capacity. Higher activity of ACS and therefore ethylene release in Varuna than RH 30 increased stomatal conductance, intercellular CO₂ concentration, carboxylation rate (carbonic anhydrase and intrinsic water use efficiency), and thus net photosynthetic rate (P _N) and leaf and plant dry masses (DM) at all sampling times. Moreover, Varuna also had larger leaf area which contributed to higher P _N and DM. A positive correlation between ACS activity and P _N and leaf area was found in both the cultivars. Thus ACS activity may affect P _N through ethylene-induced changes on foliar gas exchange and leaf growth.     

International research on cassava photosynthesis, productivity, eco-physiology, and responses to environmental stresses in the tropics

The review sums up research conducted at CIAT within a multidiscipline effort revolving around a strategy for developing improved technologies to increase and sustain cassava productivity, as well as conserving natural resources in the various eco-edaphic zones where the crop is grown, with emphasis on stressful environments. Field research has elucidated several physiological plant mechanisms underlying potentially high productivity under favourable hot-humid environments in the tropics. Most notable is cassava inherent high capacity to assimilate carbon in near optimum environments that correlates with both biological productivity and root yield across a wide range of germplasm grown in diverse environments. Cassava leaves possess elevated activities of the C₄ phosphoenolpyruvate carboxylase (PEPC) that also correlate with leaf net photosynthetic rate (P _N) in field-grown plants, indicating the importance of selection for high P _N. Under certain conditions such leaves exhibit an interesting photosynthetic C₃-C₄ intermediate behaviour which may have important implications in future selection efforts. In addition to leaf P _N, yield is correlated with seasonal mean leaf area index (i.e. leaf area duration, LAD). Under prolonged water shortages in seasonally dry and semiarid zones, the crop, once established, tolerates stress and produces reasonably well compared to other food crops (e.g. in semiarid environments with less than 700 mm of annual rain, improved cultivars can yield over 3 t ha⁻¹ oven-dried storage roots). The underlying mechanisms for such tolerance include stomatal sensitivity to atmospheric and edaphic water deficits, coupled with deep rooting capacities that prevent severe leaf dehydration, i.e. stress avoidance mechanisms, and reduced leaf canopy with reasonable photosynthesis over the leaf life span. Another stress-mitigating plant trait is the capacity to recover from stress, once water is available, by forming new leaves with even higher P _N, compared to those in nonstressed crops. Under extended stress, reductions are larger in shoot biomass than in storage root, resulting in higher harvest indices. Cassava conserves water by slowly depleting available water from deep soil layers, leading to higher seasonal crop water-use and nutrient-use efficiencies. In dry environments LAD and resistance to pests and diseases are critical for sustainable yields. In semiarid zones the crop survives but requires a second wet cycle to achieve high yields and high dry matter contents in storage roots. Selection and breeding for early bulking and for medium/short-stemmed cultivars is advantageous under semiarid conditions. When grown in cooler zones such as in tropical high altitudes and in low-land sub-tropics, leaf P _N is greatly reduced and growth is slower. Thus, the crop requires longer period for a reasonable productivity. There is a need to select and breed for more cold-tolerant genotypes. Selection of parental materials for tolerance to water stress and infertile soils has resulted in breeding improved germplasm adapted to both favourable and stressful environments. An erratum to this article is available at .     

Effect of Fruiting on Leaf Gas Exchange in Olive (Olea Europaea L.)

The effect on traits of photosynthesis and water relations of assimilate demand was studied in olive tree that has strong alternate bearing. The diurnal and seasonal leaf gas exchanges, area dry mass, and saccharide and chlorophyll (Chl) contents were measured by comparing shoots with fruit of "on-trees" (heavy fruit load) with shoots without fruit on both "on-trees" and "off-trees" (light fruit load). In spite of large seasonal and diurnal differences, leaf net photosynthetic rate (P _N), stomatal conductance (g _s), sub-stomatal CO₂ concentration (C ₁), transpiration rate (E), and respiration rate (R _D) were not significantly influenced by fruit load or by the presence or absence of fruit on the shoot. An only exception was at the beginning of July when the one-year-old leaves on shoots with fruit had slightly higher P _N and E than leaves on shoots without fruit. Water content, Chl and saccharide contents, and area dry mass of the leaf were not substantially influenced by the presence/absence of fruit on the shoot or fruit load. Hence the sink demand, associated with fruit growth, did not improve leaf photosynthetic efficiency in olive. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of reduced irradiance on leaf morphology,photosynthetic capacity,and fruit yield in olive (<Emphasis Type="Italic">Olea europaea</Emphasis> L.)

One-year-old olive trees (cv. Koroneiki) were grown in plastic containers of 50 000 cm³ under full daylight and 30, 60, and 90 % shade for two years. The effects of shade on leaf morphology and anatomy, including stomatal density and chloroplast structure, net photosynthetic rate (P _N), stomatal conductance (g _s), and fruit yield were studied. Shade reduced leaf thickness due to the presence of only 1–2 palisade layers and reduced the length of palisade cells and spongy parenchyma. The number of thylakoids in grana as well as in stroma increased as shade increased, while the number of plastoglobuli decreased in proportion to the reduced photosynthetically active radiation (PAR). The higher the level of shade, the lower the stomatal and trichome density, leaf mass per area (ALM), g _s, and P _N. Shade of 30, 60, and 90 % reduced stomatal density by 7, 16, and 27 %, respectively, while the corresponding reduction in P _N was 21, 35, and 67 %. In contrast, chlorophyll a+b per fresh mass, and leaf width, length, and particularly area increased under the same shade levels (by 16, 33, and 81 % in leaf area). P _N reduction was due both to a decrease in PAR and to the morphological changes in leaves. The effect of shade was more severe on fruit yield per tree (32, 67, and 84 %) than on P _N indicating an effect on bud differentiation and fruit set. The olive tree adapts well to shade compared with other fruit trees by a small reduction in stomatal and trichome density, palisade parenchyma, and a significant increase in leaf area.     

Relationship between root vigour,photosynthesis and biomass in soybean cultivars during 87 years of genetic improvement in the northern China

During the last century, the world soybean yield has been constantly enhancing at a remarkable rate. Factors limiting the soybean yield may be multiple. It is widely acknowledged that changes of root metabolism can influence aboveground characteristics, such as the seed yield and photosynthesis. In this study, we considered root bleeding sap mass (BSM) and root activity (RA) as indicators of the root growth vigour. We used 27 soybean cultivars, spanning from 1923 to 2009, to evaluate the contribution of root characteristic improvement to efficient photosynthesis and dry matter production. The BSM, RA, net photosynthetic rate (P_N), and organ biomass were measured at different growth stages, such as the fourth leaf node, flowering, podding, and seed-filling stage. Our results showed that the soybean cultivars increased their biomass and P_N thanks to genetic improvement. At the same time, BSM and RA also increased in dependence on a year of cultivar release. However, both P_N and biomass were positively correlated with root characteristics only at the podding stage. Our data revealed that the improved root characteristic may have contributed to the enhanced photosynthesis, biomass, and yield of soybean cultivars during last 87 years of genetic improvement. We suggest that BSM and RA could be used as important indexes for further practice in soybean production improvement.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Comparison of the Drought Stress Responses of Tolerant and Sensitive Wheat Cultivars During Grain Filling: Changes in Flag Leaf Photosynthetic Activity,ABA Levels,and Grain Yield

Water status parameters, flag leaf photosynthetic activity, abscisic acid (ABA) levels, grain yield, and storage protein contents were investigated in two drought-tolerant (Triticum aestivum L. cv. MV Emese and cv. Plainsman V) and two drought-sensitive (cvs. GK élet and Cappelle Desprez) wheat genotypes subjected to soil water deficit during grain filling to characterize physiological traits related to yield. The leaf water potential decreased earlier and at a higher rate in the sensitive than in the tolerant cultivars. The net CO₂ assimilation rate (P _N) in flag leaves during water deficit did not display a strict correlation with the drought sensitivity of the genotypes. The photosynthetic activity terminated earliest in the tolerant cv. Emese, and the senescence of flag leaves lasted 7 days longer in the sensitive Cappelle Desprez. Soil drought did not induce characteristic differences between sensitive and tolerant cultivars in chlorophyll a fluorescence parameters of flag leaves during post-anthesis. Changes in the effective quantum yield of PSII (Φ_PSII) and the photochemical quenching (qP) depended on the genotypes and not on the sensitivity of cultivars. In contrast, the levels of ABA in the kernels displayed typical fluctuations in the tolerant and in the sensitive cultivars. Tolerant genotypes exhibited an early maximum in the grain ABA content during drought and the sensitive cultivars maintained high ABA levels in the later stages of grain filling. In contrast with other genotypes, the grain number per ear did not decrease in Plainsman and the gliadin/glutenin ratio was higher than in the control in Emese during drought stress. A possible causal relationship between high ABA levels in the kernels during late stages of grain filling and a decreased grain yield was found in the sensitive cultivars during drought stress.     

Effects of Long-Term Fertilization on Leaf Photosynthetic Characteristics and Grain Yield in Winter Wheat

Based on a 20-year fertilization experiment with wheat-maize double cropping system, the effects of different long-term fertilization treatments on leaf photosynthetic characteristics and grain yield in different winter wheat (Triticum aestivum L.) cultivars were studied in the growing seasons of 2000–2001 and 2001–2002. A total of nine fertilization treatments were implemented, i.e. no fertilizer (CK), N fertilizer (N), N and P fertilizers (NP), N and K fertilizers (NK), N, P, and K fertilizers (NPK), only organic manure (M), organic manure and N fertilizer (MN), organic manure and N and P fertilizers (MNP), and organic manure and N, P, and K fertilizers (MNPK). With the treatments of combined organic manure and inorganic fertilizers (TMI), net photosynthetic rate (P _N), maximal activity of photosystem 2, PS2 (F_v/F_m), and chlorophyll content (SPAD value) of flag leaves and leaf area index (LAI) were much higher at the mid grain filling stage (20 or 23 d post anthesis, DPA), and exhibited slower declines at the late grain filling stage (30 DPA), compared with the treatments of only inorganic fertilizers (TI). The maximal canopy photosynthetic traits expressed as P _N×LAI and SPAD×LAI at the mid grain filling stage were also higher in TMI than those in TI, which resulted in different grain yields in TMI and TI. Among the treatments of TMI or among the treatments of TI, both flag leaf and canopy photosynthetic abilities and yield levels increased with the supplement of inorganic nutrients (N, P, and K fertilizers), except for the treatment of NK. Under NK, soil contents of N and K increased while that of P decreased. Hence the unbalanced nutrients in soil from the improper input of nutrients in NK treatment were probably responsible for the reduced flag leaf and canopy photosynthetic characteristics and LAI, and for the fast declining of flag leaf photosynthetic traits during grain filling, resulting in the reduced yield of NK similar to the level of CK.     

Canopy and leaf photosynthetic characteristics and water use efficiency of sweet sorghum under drought stress

The objective of this paper was to examine the relationship between Water Use Efficiency (WUE) at the canopy and leaf levels, to determine soil moisture conditions, which can optimize yield, and WUE of sweet sorghum (Sorghum bicolor (Linn.) Moench), thus providing some theoretical foundation for using marginal land effectively and developing production of sweet sorghum. Three levels of soil moisture conditions were established, and photosynthetic characteristics and yield were measured. The canopy apparent photo-synthetic rate (CAP) and leaf photosynthetic rate (P _N) were reduced gradually with increased drought stress, and the CAP was lower than the P _N under every soil moisture conditions. The P _N had a midday depression phenomenon, but the CAP did not exhibit this midday depression phenomenon under severe drought stress. The linear regression relationship of CAP and P _N was CAP = 1.5945 + 0.1496 P _N. The canopy apparent WUE_C and leaf WUE_L were the highest under moderate drought stress. The first was 5.3 and 5.8 times higher than the WUE_L in mid-July and late August, respectively. The stem fresh biomass yield was 77 tons/ha under moderate drought stress and WUE of aboveground biomass yield (WUE_B) was also the highest. Our results showed that moderate drought stress did not result in a significant reduction in biomass yield but increased WUE significantly.     

Photosynthesis,Transpiration, and Water Use Efficiency of Vegetative and Reproductive Shoots of Grassland Species from North-Eastern China

The differences in net photosynthetic rate (P _N), transpiration rate (E), and water use efficiency (WUE) between the vegetative and reproductive shoots of three native grass species from the grassland of northeastern China [grey-green and yellow green populations of Leymus chinensis (Trin.) Tzvel., Puccinellia tenuiflora (Griseb) Scrib & Merr, Puccinellia chinampoensis Ohwi] were compared. The two type shoots experienced similar habitats, but differed in leaf life-span and leaf area. The leaf P _N and WUE for the vegetative shoots were significantly higher than those for the reproductive shoots in the grasses, while their E were remarked lower in the dry season. Relative lower leaf P _N and WUE for the reproductive shoots of grassland grasses may explain the facts of lower seed production and the subordinate role of seed in the grassland renewal in north-eastern China.     

The Cause of the Difference in Leaf Net Photosynthetic Rate between Two Soybean Cultivars

To explore the cause of difference in photosynthetic performance between different cultivars of crops, leaf net photosynt rate (P _N) and photosystem 2 (PS2) photochemical efficiency (F_v/F_m), apparent quantum yield of carbon assimilation (_c), electron transport rate, photophosphorylation activity, etc. were measured in two soybean cultivars, Heinong 42 and Heinong 37. At pod setting and filling, significant differences in P _N between them were observed. The former with a higher P _N (from 7 to 38 %) had a significantly higher leaf thickness, leaf dry mass/area (LMA), chlorophyll content, soluble protein content, apparent quantum yield of electron transport through PS2 (_e), carboxylation efficiency (CE), and ribulose-1,5-bisphosphate carboxylase (RuBPC) activity. The significantly higher P _N of Heinong 42 is mainly due to its higher content and activity of RuBPC.     

Effects of Angular Leaf Spot and Rust on Leaf Gas Exchange and Yield of Common Bean (Phaseolus Vulgaris)

Isolated and interactive effects of angular leaf spot (caused by Phaeoisariopsis griseola) and rust (caused by Uromyces appendiculatus) on leaf gas exchange and yield was studied in common bean (Phaseolus vulgaris L. cv. Carioca) plants. Gas exchange was measured on 37, 44, 51, and 58 d after planting using a portable photosynthesis system. The inoculation of plants with P. griseola (P), U. appendiculatus (U), and the combination of both pathogens (P+U) caused a significant reduction of net photosynthetic rate (P _N) and yield. The reduction of stomatal conductance (g _s), P _N, and yield was higher under P and combination of P+U than under U treatment. By effect of U, the reduction on yield was higher than the reductions on gas exchange parameters. On the treatment P+U, a reduction of 23 % in P _N and a correspondent reduction of 32 % in yield was observed. The interactive effects of the pathogens on yield could be explained in part by the decreases in g _s and in P _N of diseased bean leaves. The combined effect of both diseases on yield and gas exchange parameters suggests an antagonistic interaction.     

Genotypic variation for drought tolerance in cotton (Gossypium hirsutum L.): Leaf gas exchange and productivity

Although water-limited environments are detrimental to cotton growth and productivity worldwide, identification of cotton (Gossypium hirsutum L.) genotypes that are less sensitive to drought may improve productivity in drought prone areas. The objective of the study was to assess genotypic variation for drought tolerance in cotton varieties using physiological attributes as selection criteria, and to determine the relationship of physiological attributes with productivity traits. The association of target physiological traits for drought tolerance (photosynthetic rate (P_n), stomatal conductance (g_s), and transpiration rate (E)) with productivity traits under well-watered (W₁) and water-limited (W₂) regimes was analyzed using 32 public cotton cultivars/bred lines in two field experiments conducted during the normal cotton growing seasons 2003 and 2004. Seed cotton yield (SCY) and biological yield (BY) were markedly affected under W₂ regime in all cultivars except the outstanding performance of CIM-1100 and RH-510 proving their superiority to other cultivars in drought tolerance. Conversely, FH-901, FH-634, and FH-2000 were high yielding under W₁ regime; however, exhibited a sharp decline in yield under W₂ regime. A positive correlation between SCY and BY under water stress (r=0.44 in 2003; r=0.69 in 2004) indicates that BY is also a primary determinant of SCY under water stress and genetic improvement of BY under water-limited environment may also improve SCY. P_n, g_s, and E were significantly reduced by water stress. Substantial genotypic variation for gas exchange attributes existed among the cotton cultivars. A positive association (P<0.01) was observed between g_s and E under both regimes in both years indicating the prevalence of stomatal control of transpiration. The positive association (P<0.01) between P_n and g_s in both years in W₂ regime indicates also a major role of stomatal effects in regulating leaf photosynthesis under water-limited conditions. P_n was significantly correlated with SCY (P<0.01) and BY (P<0.05 in 2003; P<0.01 in 2004) in W₂ regime; however, the level of these associations was not significant in W₁ regime. These findings demonstrate that association of P_n with productivity is effective under water-limited environment and may be useful as a selection criterion in breeding programs with the objective of improving drought tolerance and SCY under water-limited environments. Moreover, association between SCY and BY under water stress suggests that genetic improvement of BY under water stress may also improve SCY.     

Photosynthesis,Transpiration, and Water Use Efficiency of Two Puccinellia Species on the Songnen Grassland,Northeastern China

The intra- and inter-specific variations in net photosynthetic (P _N) and transpiration (E) rates and water use efficiency (WUE) of Puccinellia tenuiflora and Puccinellia chinampoensis leaves were compared. The two species experienced a similar habitat, but differed in leaf area, leaf colour, and nitrogen contents. Leaf P _N and E for both reproductive and vegetative shoots of the two species declined with leaf age. P _N for reproductive shoots was less than for vegetative shoots, but their E was greater than that of vegetative shoots in the dry season. The average P _N and E for reproductive shoots of P. tenuiflora were lower than those of P. chinampoensis, but higher for vegetative shoots.